Hikobial3：645-665.2002
Molecularphylo窪enyOfhypnobrJ/aleanmOssesasin化rredfroma lar淫e-scaledatasetofchlOroplastlｂｃＬ，withspecialre他rencetothe HypnaceaeandpOssiblyrelatedfnmilies1
HIRoMITsuBoTA，ＴｏＭｏＴｓｕＧｕＡＲＩＫＡｗＡ，ＨＩＲｏＹｕＫＩＡＫＩＹＡＭＡ，ＥＦＲＡＩＮＤＥＬｕＮＡ，DoLoREs GoNzALEz,MASANoBuHIGucHIANDHIRoNoRIDEGucHI
TsuBoTA,Ｈ､,ARIKAwA,Ｔ,AKIYAMA,Ｈ,,ＤＥＬｕＮＡ,Ｅ,GoNzALEz,，.,HIGucHI,Ｍ ４ ＆DEGucHI，Ｈ、2002.Ｍolecularphylogenyofhypnobryaleanmossesasinferred fiPomalarge-scaledatasetofchloroplastr6cL,ｗｉｔｈspecialreferencetotheHypnaceae andpossiblyrelatedfamiliesl3:645-665．
▲ Phylogeneticrelationshipswithinthehypnobryaleanmosses(ｉｅ,theHypnales,Leuco- dontales,andHookeriales）havebeenthefbcusofmuchattentioninrecentyears Herewepresentphylogeneticinfierencesonthislargeclade，andespeciallyonthe Hypnaceaeandpossiblyrelatedftlmilies，basedonmaximumlikelihoodanalysisof l81r6cLsequences、Oursmdycorroboratesthat（１）theHypnales(ｓｓｔｒ.[＝sensu Vittl984])andLeucodontalesareeachnotmonophyleticentities、TheHypnalesand LeucodontalestogethercompriseawellsupportedsistercladetotheHookeriales;(2) theSematophyllaceae(s」at［＝sensuTsubotaetaL2000,2001a,ｂ])andPlagiothecia‐ ceae(s・str.［＝sensupresentDareeachresolvedasmonophyleticgroups,whileno particularcladeaccommodatesallmembersoftheHypnaceaeandCryphaeaceae;ａｎｄ (3)theHypnaceaeaswellasitstypegenusﾉﾘDlwz"川tselfwerepolyphyletioThese resultsdonotconcurwiththesystemsofVitt(1984)andBuckandVitt(1986),who suggestedthatthegroupswithasinglecostawouldhavedivergedfiFomthehypnalean ancestoratanearlyevolutionarystage，fbllowedbythegroupswithadoublecosta (seealsoTsubotaetall999;Bucketal2000）OurresultsfiPomlikelihoodanalyses ofalarger6cLdatasetconfinnthesuggestionbysmallerdatasetsfiommultipleloci firstbyDeLunaetal.(2000)andthenBucketaL(2000)thatthereisnoevidencefbr supportingtheHypnales(ｓｓｔｒ.[＝sensuVittl984])andLeucodontales・Adetailed reconsiderationofordinalclassihcationandfnmilialclassificationwithintheHyp- nidaeiｓｎｏｗnecessary．
KeyWords:HypnalesLeucodontales,Hookeriales,hypnobryaleanmosses,molecular phylogeny,r6cLlargedataset,Hypnaceae,Sematophyllaceae,Plagiotheciaceae,Am- blystegiaceae
。 肋Ｄ'"ｊ７Ｍ６ｏ/ｑｄＭｒﾉﾉ℃"ＭＤｅｇ"ｃｈｊ,Depα""ze"ｔｑ/Ｂｊｏ/ogjcα/Ｓｂｊｅ"Ｃｅ,Ｇｒａｄ"α/e Sb/ioo/ｑ／血e"Ｃｅ,Ⅲ℃sﾉｶ〃α〔ﾉﾉｊｊｖｅｌＭ１ﾉ,hgamli〕ﾉα碗αノー3-/,〃jgZJSﾙﾉｰﾊjrosﾉi〃α- Ｗ,ﾉﾌﾞIﾉ･Csﾉi〃α739-852．ﾉﾋＷ/L 7bmojiFz`9ＭﾉWmwq,Depαﾉﾌﾞ'"e"'０/Ｂｊｏ/Ogjcα/Sbje"Ｃ＆Ｆ,Ｇ７α`"α/e此/ＩＯＣ/q/ｍｉｅ"Ｃｅ， U>ﾌﾙe'Ｍﾜﾉq/mtyo,〃O'7907-3-/,βz"7/bﾉo-k"’７bﾉb′o川-0033,上pα"・ 航rqMEMﾉhi〕′α',、,/Ｍｌｓａｌ''１ｑ／ﾉVb【/"'℃α'z`ﾉ仙加α'Ｍα〃jrjes,町ogo，YZlO′ojgaoAa 6-cho"ze,５位"dtJ-sAj,Hyogo669-/ｺﾞｲｄﾉﾋZpα〃 〃αｍＤｅＬ""α＆ＤＣ/oresOo'1Ｚ｡/ez,ＤｅＰａｒｔａｍｅ"ｊＭｅＳＭ伽/ｊｃａﾘﾉｾge/αＭＭｍｔｏ ｃ/ｅＥｃｏ/ｏｇｍｌＡＣ.,ＸＭＪｐａ，比７．９/000,ＭｄＸｊｃｏ、 MJSα"０６〃ｆﾉﾉgZlcAj，Depα"ｍｅ"／ｑ/ＤＣ/αノリＭＪ/jolZα/比ｊｅ"ｃｅＭ`Ｓｃ"〃，伽αAr"６０ ｲｰ/-ＩｍＭ"6α-sﾙﾉ,仏aMhH305-000LﾉﾋZpα",ＤＥＰα"ﾉ?Te"／ｑ/Ｂｉo/ogjcα/Ｓ℃/e"ＣＧＳ， G/ndZJα/ｅ比ﾉioo/ｑ/此ｊｅ"Ce,Ｕ>7〃e′Ｍﾜﾉｑ/、oﾉb′０，肋"ｇｏ７~3-/,Ｂ２`"ﾉtyo-ﾉ1Ｗ,Tbﾉbﾉ。 〃3-0033,上pα〃 'ContributionfiOmtheLaboratolyofPlantTaxonomyandEcology,HiroshimaUniversity・ＮＳｅｒ・No.S２Ｌ 646 HikobiaVoll3,No.４，２００２
Introduction talesandHookeriales）fbrcorroborationofthe phylogenetichypothesisofDeLunaetal.(2000） Thehypnobryaleanmosses，includingtheor- ａｎｄＢｕｃｋｅｔａＬ(2000);３)toclarifythefiamilial dersHypnales，LeucodontalesandHookeriales， relationships，especiallybetweentheHypnaceae areoneofthemostdiversifiedmossgroups、 andrelatedfamilies，suchastheSematophylla- Attentionhasbeenpaidtotheclassiflcationof ceaeandPlagiotheciaceae，withinthehypnobly- thehypnobryaleanmossessinceVitt（1984)pro‐ aleanmosses，giventhattheLeucodontalesand videdanewsystemfbrtheBryopsida、Within Hypnalesarenotmonophyletic;ａｎｄ４)totracein thepleurocarpshesuggestedthat“adoublecosta thephylogenythetransitionsofthecostafiDm ll groupasrepresentedbytheHypnaceae,Selnato- singletodouble，oneoftheimportantcharacters phyllaceaeandEntodontaceae，isdivergentfiDm fbrthefiamilialclassificationofthehypnobry- ０ agroupwithasinglecosta，ｗｈｉｃｈｃａｎｂｅｒｅ－ aleanmosses、 ｇａｒｄｅｄａｓｔｂｅｍｏｓｔｒecentevolutionarystageof Recent，molecularphylogeneticanalysesfbr theorderHypnales(seealsoTsubotaetall999； pleurocarpousmosseswerebasedonr6cLse- Bucketal2000）Subsequently,BuckandVitt quencesalongwithotherdatasets,ｓｕｃｈａｓａｍＢ， ￣ (1986)constluctedanotherclassificationfbrhyp- 伽Ｌ－Ｆａｎｄノアs４Ａｔｔｈｅｐｒｅｓｅｎｔｔｉｍｅ，tbere- noblyaleanmossesbasedonmorphologicaland sourcesofr6cLsequencesarethemostcompre- ecologicaldata、 hensivesamplingfbrthehypnaleanmosses・ Inrecentyears，phylogeneticrelationships Alignmentofthe'6cLsequencesisalsoeasybe- withinthehypnobryaleanmosseshavebeenthe causeoftheirdistinctionasaprotein-codingre‐ fbcusofmucbattentionatseveraltaxonomiclev- gioninthecodonpositions・Sqwepresenthere els(Arikawa＆Higuchil999,2002;BucketaL pbylogeneticin化rencesfbrthethreeordersHyp- 2000；ＤｅＬｕｎａ＆coworkersl999，2000；Maeda nales，LeucodontalesandHookeriales，basedon etaL200qTsubota&coworkersl999,2000, themaximum-likelihood（ＭＬ）analysesofl81 2001a,ｂ;VanderpoortenetaL2002)．Tberela- sequencesofthechloroplastribulosel,5-bisphos- tionshipswithinthehypnobryaleanmosseshave phatecarboxylase/oxygenaselargesubunit beenevaluatedbybothmorphologyandmolecu- (r6cL),togetherwithrecentinfbnnationonthe larstudies(Hedeniisl995;ＤｅＬｕｎａｅｔａＬ１９９９)． molecularphylogenyofhypnobryaleanmosses， Hedeniis（1995,1996)showedthemonophylyof anddiscusstheimplicationsofourfindingson theHookeriales-Sematophyllaceaeanddiscov- theHypnaceaeandtheirrelatedfamilies． eredtheLeucodontalesasagradetaxonbasedon morphologica1data、DeLunaetaL(1999)clari- Materialsandmethods fledthehomologyofpleurocarpyandthemono- phylyofallhypnobryaleanpleurocarpousmosses Phylogeneticanalysesbasedonther6cLgene (theLeucodontales,HypnalesandHookeriales)， sequenceswerebasicallyperfbnnedfbllowing basedonacombined'6cLsequencesandmor- Tsubotaandcoworkers（1999,2000,2001a,ｂ)． phologydataset・Furtherdetailswererecently Themethodconsistsoftwosteps：Dobtaining の revealedbyDeLunaetaL(2000)andBucketal． sequencedata(DNAextractionPCRamplifica- (2000)whoproposednewphylogeneticrelation- tionandDNAsequencing,orDNAdatabaseho- shipswithinthesepleurocarpsTheseworks mologysearch);and２)dataanalyses． suggestedthattheLeucodontalesandHypnalesas previouslyunderstoodarenotmonophyletic ８℃"7calα"c/Ｐ/α"ｒＭＪＺｅｒｊａん Familiesfroｍｔｈｅｓｅｔｗｏｏｒｄｅｒｓａｒｅｍｉｘｅｄｉｎａ Atotalofl81mossspecieswereexaminedfbr ｓｉｎｇlelargecladesistertothehookerialeanlin- theanalysesasshowninTablelandinAppendix eage、 A・Thelattercontainsvoucherinfbmationand Theobjectivesofthisstudyare:ｌ)toconstruct DNADatabaseAccessionNos・Voucherspeci- aphylogenetictreefbrhypnobryaleanmosses menssequencedinthepresentsmdyweredepos- basedonlarge-scaledataset;２)totestthemono- itedi、TNS,ＨＹｏ,orHIRoSomesequencesused phylyofthethreeorders（Hypnales,Leucodon- fbrtheanalyseswerealsoobtainedfi･omthe DNAdatabase． HIsuBoTA,Ｔ・ARIKAwA,Ｈ・AKlYAMA,ＥＤＥＬｕＮＡ,Ｄ､OoNzALEz,MHIGucHIANDH､DEGucHI ６４７
TableLListofspeciesandftlmiliesinvestigatedlbrl6cLgenewiththeaccessionnumberandvoucheror referenceofthesequences、TreatmentoffamiliesinthistablefbllowsVitt（1984)ｗｉｔｈsomemodificationsby BuckandVitt(1986)．FurtherinfbnnationfbrthesequencesobtainedinthepresentsmdyisshowninAppendix Ａ、
OrdelﾌﾟFamiIv/SDccies AccessionNo． Refcrcnccs/Origin HypnaleS(s・stⅨ） Hypnodendraceac A0lp"ocノセ"cjWmel2zjesjj(Hook.)Paris ＡＦ２３ＩＯ９３ MishIcrctal(onIyinDNA database） PIeuroziopsidaceae 雄"mzjqpsな'wﾉﾉze"たα(Weinm.)Kindb・cxBritt、 ＡＢＯ24683 ArikawaamdHiguchi(19”） Fontinalaceac Fo"""α/jFampW℃"ｃｑＨｅｄｗ ABO50949 msubotaetaL(2001a） ｡ Ｆｂ"'ｊ"αﾉ応α"伽ﾉｱ'eZjcqHedw、 AJ275183 Coxctal(2000） Ｆｂ"/、αﾉﾉＳ“んcar/ｊｃａＢｒｕｃｈ＆Schimp・ AF231074 DeLunaetaL(2000） Echinodiaceae Ebﾉｸmodmll2〃"61℃s"'?z(Mitt.)Ａ・JaCgcr AF233568 DcLunaetal(2000） Sbmrommm〃jcosjcJm”(SulL)Mitt、 ABO24677 Arikawa＆Higuchi(1999） Thuidiaceae j6jerj"eﾉﾉﾋJa6jerj"α(Hedw.)Ｍ・Fleisch ＡＦＯＯ５５１９ Goffinctctal(1998） ７１ﾉｶ､とﾉﾉ"ｍｃﾉｾﾉｊｃａｍｍｍ(Ｈｅｄｗ)Schimp、 ＡＦＩ５８１７７ DcLunactaL(2000） ７１ｌｂ､ﾋﾉmmplな'ocaOMMUⅡ､HaL)AJacgcr ＡＢＯ７１４Ｉ６ TSubotactal(200ｌｂ） nh"､１１"〃だcqg"""伽(Hedw.)Lindb． ＡＢＯ１９476 Akiyamactal.(2000;only inDNAdatabascXmsubota etal(2001a） Lcskcaceae Bom〃αｍｊ"e"ﾉﾉ(Broth.)Ｃａｒｄｏｔ ＡＢＯ２４９６３ TBubotactal(1999） Ｄｍ/tieﾉﾉﾋJSpecjosjFsjmqBrothexCardot ＡＢＯｌ９４６７ Akiyamaetal.(2000;only inDNAdatabasc);thiSstudy M〕′αbeαﾉﾘw"ＣｅﾉﾉﾋJ(Mitt.)Broth ＡＢＯｌ９４７５ Maedaetal(2000） OAcJm皿池ｅａｈａＡｏ"je"sjs(Mitt.)Broth ＡＢＯＩ９４７７ Macdactal(2000） Amblystegiaceac CtJ"je堰o"eﾉﾉﾋJc"sp此ｍね(Hedw.)Loeske ABO24678 Arikawa＆Higuchi(1999） C'、o"e"'℃"〃ｃｊ'､"(Hedw)Spruce ABO95270 thisstudy D'℃pα"ocmdi`sadi`"c"s(Hcdw.)Warnst． ABO24681 Arikawa＆Higuchi(1999） ｆbｲ91℃α''161ｌＷｅｇｊ"〃'e'､x(Hedw.)Jcnn AF23356５ DeLunactal.(２０００） Lembophyllaceae Doﾉﾉcﾉｶo､"rjqpsjM1ve応施'ｍな(Mitt.)Ｎｏｇ ABOI9465 Akiyamactal(2000;only inDNAdatabasc);thisstudy Ｌｅ"16qpﾉＭ/""'cﾉﾙ"/Sm"?(Hookf＆Wilson)Lindb， AF233570 DeLunactal.(2000） StcreophyIlaceac EmodO"'qpsハノＢＨＣＯＳ／２gα(Brid.)ＷＲＢｕｃｋ＆lreland ABO24635 Arikawa＆Higuchi(1999） S1e花qpﾉ!)ﾉﾉﾉ"加沢adIczjﾉCs"、(Hook)MitL ABO24637 Arikawa＆Higuchi(1999） Fabroniaceae A"αcα'Modb"spmch"o雄s(FroeLexBrid.)Ｂｒｉｄ AF231077 DeLunaetal.(2000） 〃ﾋﾞﾉﾉcodO"""maJpjﾉﾉﾋJ”(Hcdw)AJaeger AF23357］ DeLunaetal.(2000） Brachythcciaccac Bmc〃んecj"ｍｐ肋､Cs"、(Hedw.)Schimp、 AF233566 DeLunactaL(2000） Ｂｍｃﾉi〕ﾉﾉﾙecj"mrjw`ﾉα”Ｓｃｈｉｍｐ ABO24674 A｢ikawa＆Higuchi(1999） Ｂｍｃ/Ｍｈｅｃｍｍｓα/e6ms"、(HofYinexRWCber＆ AFl58I76 DeLunaetal(2000） DMohr)Schimp・ ノMb'Ｈｍｃﾉａｄｔｗｃｍｍｏｗｊｃｚｊｊ(Borszcz.)Stcere＆Ｓｃｈｏｆ ABO29389 Tsubotactal.（1999） Ｐﾉﾛﾘﾉ!〕や"jdm加吻α"o/“s(Hedw.)Dix， ABO29385 Tmbotactal(1999） PseWdOsc化mpodmmpⅢ'ｗｍ(Hcdw.)Ｍ・F1cisch． AF233567 DCLunaCtal(2000） Ｒ/i)′"c/zos/egm"'pαﾉﾉﾉ｡!/bﾉﾉz`"1(Mitt.)AJacgcr ABO24944 Tsubotactal(1999） ７bme"卯""〃〃/花"s(Hedw.)Loeske ABO24676 ArikawaandHiguchi(1999） Entodontaccae E7Troc！O〃cﾉｶαﾉﾉelZgelj(Paris)Cardot ABO50993 TBubotaetal.(2001a） E)?/odb"/”ｊｍ４ｓ(Gri征)AJacgcr ABO50994 T1subotactaL(2001a） 勘/odO""!)'”“(Hook)Hampe ABO24640 Arikawa＆Higuchi(1999） EmodO"'w6jc""c/bｲs(Mitt.)AJacger＆Sauerb・ ABO29386 T1subotaetal(1999） EmodO〃sca61j血"sLindb ABO50995 TEubotaetal(200]a） Sematophyllaccae 化α"'んoW2〕''７chjzmzpqpj/、'"、(Ilarv.)Ｍ・Fleisch ＡＢＯ５１２２４ 辰msubotactal(2001a） 化mpo"zｲｍｐｚｍｇｅ"s(Hedw.)Broth ＡＦ２３３５７２ DcLunaetal(2000） jcmporm〃s"α､j"e瓦加(Reinw.＆Ｈｏｍｓｃｈ)Ｍ・Flcisch． ＡＢＯ５Ｉ２２５ T1subotactal(2001a） 648 ＨｉｋｏｂｉａＶｏＬｌ３,No.４，２００２
TableLcontinued
OrderﾌﾟFamilv/SDccies AccessionNo． Rcfercnccs/ｏｒ 、
幼ＩＤ'cﾉｶeﾉﾉZzgﾉomelmqpmpqg"ﾉ娩'α(Tbyama)Ｓｃｋｉ ＡＢＯ５Ｉ２Ｉ７ TmbotactaL(200Ia ｣5ﾉｾﾉempﾉﾘﾉﾉﾉﾉ""1〃"e(Hook.)Ｍ・F1cisch ＡＢＯ５１２１８ T3ubotaetaL(2001a ノ5leZemp/q〕ﾉﾉﾉﾉⅢ、〃e"Taros皿"ZBrothcxPLdelaVardc＆Th6m ＡＢＯ２９３９１ T1subotactaL（1999） KsocﾉUdjeﾉﾉﾋ７３脚”"ﾉﾋJrな(Dixon)ＢＣ・T1an＆Mohamcd ＡＢＯ３９７８４[asⅣｅａｃｍ‐ T1subotactal(2000） porm"Z加ｇＥﾉﾉ(/を'W"?］ Mtzsjqpo"zap皿ﾉCheﾉﾉﾋJ(Herzog)incd ABO71410[asms- Tsubotactal.(200lb） megjsljap"ﾉb〃eﾉﾉﾋJ］ MJslqpollzas皿lWZ/blwllzHorik＆Ａｎｄｏ ＡＢＯ７Ｉ４Ｉｌ T1subotactaI.(200lb） ﾉMJS'qpo"Tα皿"cj"(/bﾉ､"'(Broth)Broth ＡＢＯ７１４１２ T1subotactaI.(200lb） Mbjoﾉﾉiecj"mmJcrocα'p"、(Hook)Mitt． ＡＢＯ５Ｉ２２３ Tmbotactal(2001a） Ｐ(Jpj"jdiqpsなｍａｃｍｓＺｊａｑ(Broth.＆Paris)Ｗ,RBuck＆ ＡＢＯ５１２２０[asRhqpﾙﾉﾋﾉO‐ Tmbotactal.(2001a） Ｂ・Ｃｍｌａｎ s'ｊｃ/ｌｗｍｍａｃ'･Cs"c/"､］ 応e"ｄｏ"jm7TegjSjjaⅢ"｡iｲﾉ、α(Broth＆Ｍ・Yasuda)HAkya． ABO51229[asnjs- Tmbotactal.(2001a） ロ ＆Tsubota megjs"α皿"｡！`ﾉ、α］ DﾉﾋJな/αｄｂｌｐﾉiQco"qpm"α'α(Reimers＆Sakurai） ABO39785[ａｓＢｍ‐ T1subotactaI.(2000） ＷＲＢｕｃｋ ﾉﾉ『e”ぬＣＯ"2p/α"α`α］ ｌｌ)ﾉﾉ､､α`/blpﾉlα/2mrjej(BcschcxCardot)ＷＲＢｕｃｋ ABO39786[ａｓＢｍ‐ Tmbotactal.(2000）
ﾉﾉze”ﾉﾉα/Zmrjej］ 画 けﾉﾋ』な､｡b(りんαﾉｶe"o"〃(Duby)ＷＲＢｕｃｋ AB029167[asBro‐ mSubotaetal.（1999） /ｈｅ肥/ね/ie"o"ﾉﾉ］ ＲｙﾉﾋJjSmde Ｒﾉﾘﾉﾉﾉ“α`、ＩＤﾉmMZpo"ｊｃⅢs(Reimcrs)Ｔｌ・Ｋｏｐ ABO39788 msubotaetaL(2000） □ Ｒ/Mjdm火ｌＩｐｈ"sﾉo1℃"s(Hcdw.)Wamst． ABO24666 Arikawa＆Higuchi(1999） Ｒ/Mjdm“lIpﾉiWss9"qrmmJs(Hcdw.)Warnst． ABO24667 Arikawa＆lliguchi(1999） Ｒ/Mjdmdblph"s〃""eﾉﾉWS(Hedw)Wamst・ ABO24668 Arikawa＆HiguchiO999） Plagiothcciaccae LsqpZeﾉﾌｲgjqpsjS"ｍｅ比ｒｊα"ａ(Schimp)Ｚ・Iwats・ ＡＢｌ３４９４２ Arikawa＆lliguchi(2002） Pmgjo'Aecj"、。ｂ"'た"/、"、(Hedw)Schimp． ＡＢＯ２４６２３ ATikawa＆Higuchi(1999） Ｐ/tJgjorAecj"、加ｿ/o"ﾉﾉ(SulL)ＥＢ・Bartram ＡＢＯ２４６２５ Arikawa＆Higuchi(l”，） PﾉﾋZgjo/ﾉiec/""Te"刷りﾉ！〕ﾉﾉﾉⅢ"'(Cardot＆Th6Ⅸ)Ｚ・Iwats ＡＢO24626 Arikawa＆lliguchi(1999） P/cJgjo/Aecj冴加〃ecApem雄"ｍＢｒｕｃｈ＆Ｓｃｈｉｍｐ ABO24630 Arikawa＆Higuchi(1999） ＰﾉﾋJgjoﾉﾉiecj"ｍ〃emoraﾉe(MiU.)ＡＪａｅｇｅｒ ABO29387 msubotaCtaI.（]999） ＰﾉﾋｺgjofAecjw”〃"｡）ｲﾉα'"、(Hcdw.)Schimp・ ABO24634 Arikawa＆lliguchi(l”9） Hypnaceae Cre"域"ｍｍｏﾉﾉ皿Sc皿"2(Ｈｅｄｗ)Mitt． ABO24657 Arikawa＆Iliguchi(1999） Ｏﾉossadblpﾉmsqgα/αeBroth.＆Ｍ・Yasuda ABO50950 T1subotactaI.(2001a） OoﾉﾉﾋJ"ｍｍｇｌ"Csα(Mitt.)Broth ABO94341 thisstudy Go"α"mSp/e"ぬ"s(Iisiba)Nog． ABO94340 thisstudy 〃C吃qgjeﾉﾉﾋJpel．'･o6"s/α(Broth.)ＺＪｗａｔｓ・ ABl34944 Arikawa＆Higuchi(2002） ﾉﾌﾞＤＰ""mcz《p1℃sWbrmeHedw． ABO39674 T1Subotactal.(2000） HTsuBoTA,Ｔ・ARIKAwA,Ｈ・ＡＫⅣAMA,ＥＤＥＬｕＮＡ,Ｄ・GoNzALEz,Ｍ・HIGucHlANDH・DEGucHI６４９ TableLcontinued Order/Familv/SDecies AccessionNo． Refbrcnces/Ｏｒｉ 1， hbp"皿、/ｍｄｂｅ喧jjMitt、 ABO29390 TmbotaetaL（1999） ノカplT"”ｐ此'，Ｔａｇ/bmzeWilson ABO29384 Tmbotactal.(1999） ノh0p"""T〃jS'０－Wr雄(Broth)ＰaⅢ ABO24656 TmbotactaL（1999） ZSqpteﾚﾂ9ｍｍに"er皿加(Swb)Mitt・ AF2335６９ DcLunaetaL(2000） Lsqp/Clﾂｇｊ"碗Ｗ"eαノセＥＢ・Bartram ABO24650 Arikawa＆Higuchi(1999） Or/ﾉｶo肋ecj"ｍｒｚ(/bsce"s(Dicks・ｅｘＢｒｉｄ.)Schimp ABO50951 T1subotactal.(ZOOla） DﾉｍＳｊａｍ"にα'α(Hedwb)Schimp ABO24642[asＥ〕ﾉﾉａｊ－ Arikawa＆Higuchi(1999） ｓｊｅ"αｊ""にα/α］ PmOlgﾌﾉｒｊ皿加'"e"s(Brid.)Schimp ABO24645[ａＭ〕ﾉﾉαﾉｰ Arikawa＆Higuchi(1999） sje"αpol()ﾉα"ﾉﾉ､］ mxjp"y"""zao"DC"e"se(Besch)Ｚｊｗａts・ ＡＢＯ24648 Arikawa＆Higuchi(1999） Leucodontales Climaciaccac 、 dｊ〃αｃｍｍｃｊｂ"｡、雄s(Hedw.)FWCber＆Ｄ・Ｍｏｈｒ ＡＢＯｌ９４４２ Akiyamaetal.(200qonly inDNAdalabase);thisstudy ｄｊｍｑｃｊ"、ﾉﾋZpo"ｊｃ"mLindb ＡＢＯ１９４４３ AkiyamaetaL(2000;onIy inDNAdatabasc);thisstudy Neckcrace2e Bおselm〃"gz`/α'α(Mitt.)Broth． ABO94789 thisstudy 〃o"､"α“"ﾉｶⅢs'αrgjo"､""s(Mitt.)Dixon＆Ｐｄｅｌａ ABO94792 thisstudy Ｖａｒｄｅ 〃O"､ﾉﾉo`ノセ"鋤｡"Scα(peﾉﾉﾋﾞ/bﾉﾉ"、(Mitt.)Ｍ・FlCisch ＡＢＯ９４７８８ thissmdy ノVCcAcm”"lgcmMijll､HaI、 ＡＦｌ５８１７３ Maedactal(2000） ノVECにrqpsn〃"jZmｍ(Mitt.)Ｍ・Fleisch ＡＢＯ９４７９０ thisstudy Pm"α花ﾉﾉﾋJα"6唖α(Bosch＆SandcLac.)Ｍ､Fleisch ＡＢO94787 thisstudy ７１ｈａｍ"o6ひ"ｍｍ６ｓｅｒｊａｍｍ(Mitt・exSandeLac)ＢＣＴaｎ ABO94791［as71/mmlzo‐ thissmdy ｂひＨｍｓα"生ｊ］ Cyrtopodaceae Bescﾉbe1℃"ｊａ６１ｗｌ/b/jaHampc ＡＪ２７５１８４ CoxetaI.(2000） Besc/，e”"jaeにg[J"‘jSsjmaDuby ＡＦ２３ＩＯ９７ DCLunactal.(2000） Q′"qp"sseros"s(Hedw.)Hookf ＡＦ２３１０９６ MishlcrctaL(onlyinDNA databasc） CIyphacaccae Q)p"αｅａｓｊ"e"３曲Ｅ・BBartram ＡＢＯｌ９４５７ Macdactal(2000） qpmdm/qps町o6'"s(/b/ね(Nog.)Ｎｏｇ ＡＢＯｌ９４５８ Macdactal.(2000） Ｄｏ｡’【J/apo"jcaSandeLac ＡＢＯｌ９４４６ Maedactal.(2000） Pj/O"jcﾉｶqpsjSdb"【α'α(Mitt.)Besch ＡＢＯｌ９４6０ MacdactaL(2000） Anomodontaccac A"omodO〃α66死ＷａｍｓＭｉｔｔ． ＡＢＯｌ９４６８ Macdactal(2000） 」"omodO〃ｇｊｍ“/MUll・ＨａＬ ＡＢＯｌ９４６９ Macdactal(2000） Ａ"omodb"〃ｊ"oパHedw.)Ｌｉｎｄｂ ＡＢＯＩ９４７Ｉ Macdactal(2000） Ａ"omodb〃'wgE/〃(MiilLHaL)KeissL ＡＢＯl９４７０ Macdaetal.(2000） ﾉｦﾛp/oﾉﾛ〕′ｍｅ"ｊ"腕ﾉolZgjlTerve(Broth)Broth ＡＢＯｌ９４７２ Macdactal.(2000） 〃ZJpﾉo/ｌ〕ﾉﾉ，Telzj"'Ｗ,semb"jS'e(MiilLHaL)Broth． ＡＢＯｌ９４７３ Maedactal.(2000） 〃ど'1pe""e"、〃roccoae(Sull.＆Lesq.)Cardot ＡＢＯｌ９４７４ MacdaetaL(2000） Hedwigiaceae h'bclWjgjZzc"jα'α(Hcdw)PLBeauM AFOO55I7 GoffinctctaL(1998） ノWiacocαノア“ｐ"'p"、Sce"s(Brid)Paris AJ27SI71 Coxctal.(2000） Leucodontaceae A"""jc"、/brmosα"αＮｏｇ ＡＢＯＩ９４４５ MaedaetaM2000） 化/仰o"eaes9mmﾉﾉﾉ(Tb6r）Ｈ・Akiy・ ＡＢＯＩ９４４７ MacdactaM2000） ＦＤ“"Ｄｅｍｊα/とIpo"ｊｃａ(Besch)Paris ＡＢＯＩ９４５０ Macdactal(2000） FbハsZroemJα〃ecAeroj火sBroth ＡＢＯI９４４９ MacdactaL(2000） FblIsstmellnm〃jchollzj"ね(Hcdw)Lindb． ＡＢＯｌ９４４８ Macdactal(2000） Ｌe"ＣＯ血〃α'”v"℃"ｓＮｏｇ・ ＡＢＯｌ９４５３ Macdactal.(２０００） Ｌｅ"ＣＯ巾"/"ﾉﾋIce"s(Hedw）ＳｕｌＬ ＡＦ２３ＩＯ７５ MishlerctaL(onIyinDNA databasc） Ｌｅ"ＣＯ“〃mppo"ｉｃ"sNog． ＡＢＯｌ９４５Ｉ MaedactaL(2000） Ｌe"COC/b〃ＳＣＷ,olwTsなＢｅsch・ ＡＢＯｌ９４５２ Maedaetal(2000） Ｌｅ"codb"３ｃｍ”jdbs(Hedw.)Schwii伊 ＡＢＯ９５９８８ thisstudy Ｌｅ"codb"secW"ぬs(Harv.)Mitt・ ＡＢＯl９４５４ Macdactal.(2000） LeHcodb〃ｓｏﾉicZ〕ﾉａＡｊｅ"s応ILAkiy． ＡＢＯ１９４５５ Maedactal.(2000） Ｌe"ＣＯ“〃re"lpemrzJsH,Akiyb ＡＢＯｌ９４５６ MacdactaI.(2000） Ptychomniaccae PO’c"ｏｍ"ｉｏ〃αcjcHﾉﾋzだ(Brid)Mitt、 ＡＦ２３３５７６ DcLunaetaL(2000 Myuriaccae Emy”ｍｍｓｍｊｃⅢ、(Mitt.)ＮＣ９． ＡＢＯｌ９４６３ Akiyamaetal(ZOOO5only inDNAdatabase);thisstudy ＭｳﾉHrjⅢｍｈｏｃﾉzsje"erj(Schimp.)Kindb ＡＦ２３３５７５ DcLunactaL(２０００） 650 ＨｉｋｏｂｉａＶｏＬｌ３,No.４，２００２ TableLcontinued Order/FamiIv/SDccies AccessionNo． RcferCnccs/Origin Wardiaceae ﾚｲblz/ねﾉﾘlgl･ome〃jcaHarv.＆Hook ＡＪ２７５Ｉ７０ CoxetaL(2000） Prionodontaceae P"o"oｄｏ"火"s"s(SwbexHedw.)Mijll・Ｈａ１． ＡＦｌ５８１７４ DcLunaetaL(2000） ７ZJjwα"o6ひ"mspecJoszJmNog・ ＡＢＯ１９４６６ AkiyamactaL(2000;only inDNAdatabasc);thisstudy Ptcroblyaccae P'em6ひo〃qr6Wsc"mMitt・ ＡＢＯｌ９４６１ MacdactaI.(2000） Prem6ひo〃。b"sWmHornsch ＡＦｌ５８１７５ DcLunaetal(2000） P/ero6ひqpsjlForje"'αﾉﾉSsubspj′Ｈ２""α"e"s心(Broth)Ｎｏｇ ＡＢＯ１９４６２ Macdactal(2000） ｍＪｃﾉＭｏｍａｊ"ぬczJmMitt・ ＡＢＯ１９４6４ AkiyamactaI.(Z00qonIy inDNAdatabasc);thissmdy Trachypodaceaｅ ７》zzc/1)podbps応α皿"cヅノ､α(Mitt)Ｍ・F1cisch ABO24682 Arikawa＆lliguchi(]999） □ ７》zJcﾉＷ,Ⅸｓ６ｊｃｏﾉorRCinw.＆Homsch AF233577 DeLunactaL(2000） Mcteoriaccac P[Zpj"αrjZJcノピppej(HornschexMUlLHaI.)AJaeger ＡＦｌ５８Ｉ７２ DeLunaetal.(2000） Hookeriales Daltoniaccac qＰ Lepj［わ〃/ｍｍｓ"ｒｊ"αme"seMijⅡ､Hal､ ＡＦ２３３５７８ DcLunactal(2000） Hookeriaceae 〃OoAerjaacm/bﾉﾉαＨｏｏｋ＆Orev、 ＡＦｌ５８１７０ DcLunactaI.(2000） HypoptcD/giaccac ノbpqpje奴gjHmZamarなcj(Sw.)Ｂ｢id・exMUlI・Hal、 ＡＦｌ５８１７１ DcLunactal(2000） ハ0ｨpqp'Cl〕ｌｇｊ"〃/α/zj/e"seAngstrOm ＡＦ２３１０９５ MishleretaI.(onIyinDNA databasc） BryaIes BIyaceae MJeﾉﾉcﾉzAq/brjaeﾉolZgzJ'α(Hoppc＆Ｈｏｍｓｃｈ)Nccs＆ AF232693 CoxctaI.(2000） Hornsch O〃ﾉｍｄｂ"""〃ﾉj"eα”Schwiigr・ ＡＪ２７５Ｉ７４ CoxctaL(2000） ＰＣｶﾉmclwcﾉﾋJ(Hedw.)Lindb ＡＪ２７５１７５ Coxctal(2000） Lcptostomataccac LepmsjomＷｍｍａｃｍｃα'1p"、(Hcdw.)Bach・Pyl、 Ａｊ２７５１７８ CoxctaL(2000） Mniaccac A化j""ﾉﾉjomso"ｊｊＳｃｈｉｍｐ AFOO55I8 OofYinclctaI.（1998） Ｐｍｇｊｏｍ"ｊ"、cⅢSpj血/"ｍ(I-ledw.)1XLKop・ U87082lasM"ｊ"m Lewisctal（1997） cWqpjdbm'?？］ Ｐｍｇｊｏｍ"ﾉﾂﾞﾉ，，/ZJpo"jc"、(Lindb)1Ｊ・Kop・ ＡＢＯ50992 T1subotactaL(2001a） Rhizogoniaccac Bw･ﾉio6〃""2Ｖαﾉﾉﾊｰｇｍｒｍｅ(HampccxMUlLHaL)Manucl ＡＪ２７５１７９ CoxetaI.(2000） Bartramiaccac Bar"αｍｊａＰｏｌｍ/brmjSHedW ABO24620 Arikawa＆Higuchi(1999） Lejomeﾉｨz6ar"αmjo此尤s(Hook.)Ｐａｒｉｓ AF478238 Magombo(2002） Ｐﾉﾉﾉﾉo"o油α"ぬ"α(Mitt.)Ａ・Jacgcr AF478240 Magombo(2002） Aulacomniaccac A"ﾉacom"j皿、！zｨ垣竝《"(WahIcnb)Schwiigr・ ＡＪ２７５１８０ Coxctal(2000） RacopiIaceac Racqpj/ｌＪｍｃｏｍ'０ﾉⅢ/αcezｨ"(ＭＵⅡ､llaL)Rcichardt ＡＦ２３１０９４ Mishlcrctal(onlyinDNA database） Outgrouptaxa 、 Erpodiaccac I'b"'2J"eﾉﾉﾋJsj"elTsLs(Vent.)MUllHal ＡＦＯＯ５５４６ GofYinctctal（1998） SpIachnaccac ７h〕ﾉﾉoljα〃"９２`/α/α(Dicks.)Lindb ＡＦＯＯ５５１５ GofTinctctal.（1998） Orthotrichaccac D'ｗｍｍｏ"`地０６'2Ｗｂ/jaMUllllaL AF232697 Ｃｏｘｃｔａｌ(2000） ノＭｒ"ｍｊ"j"Ｗ"c"rv ＤＭ欧伽crjo",ＰＣＲ。〃/抗cαﾉﾉo〃α"c/DM aligned,thosebelongingtothesamesequenceare Sb9"e"cmg representedbyasinglesequenＣｅ，andanyun- TotalDNAiswereextractedfi･omfieshsam- alignablesequencewasomittedfiomthedata plesorherbariumspecimensbymodificationsof sets． theCTABmethod(Murray＆Thompsonl980； Arikawa＆Higuchil999;TsubotaetaL2000)or A"αbﾉsjsjWAoqls phenol-chlorofbrmmethod(TsubotaetaL1999） Treeswereconstructedbythefbllowingthree PCRamplificationsof76cLgenesegmentsand methods:theneighborjoining(NJ)(Saitou＆Nei directseqUenceanalyseswerecarriedoutusing l987)，themaximum-parsimony（ＭＰ）method standardconditionsaccordingtothemanufactur- (Fitchl971),andthemaximum-likelihood(ＭＬ） ersinstructions、Thereactionswereperfbrmedin method(Felsensteinl981).Thetreesobtainedby thethermalcyclerswithPCRamplificationkits thethreemethodswereappraisedbythelog-like- andsyntheticprimers・Thesequenceswereelec- lihoodmeasurewithNucMLiｎＭＯＬＰＨＹ２３ｂ３ trophoresedonautomatedsequencers，andana- (Adachi＆Hasegawal996)． 、伽α"ｃｅａＭ〕ﾉsjs--ANJtreewasconstructed 。 lyzedonthedata-analysissystemThesequences obtainedinthepresentstudyweresubmittedto byNuｃＭＬａｎｄＮＪｄｉｓｔｉｎＭＯＬＰＨＹｖｅｒｓｉｏn ｔｈｅＤＤＢＪ/ＥＭＢＬ/GenBanklntemationalNucle- 23b3package（Adachi＆Hasegawal996,see otideSequenceDatabaseCollaboration AppendixB)usingHKY85model(Hasegawaet DetailsandiilrtherinfbrmationfbrDNAex- aL1985)fiorthedistanceestimations． tractingprotocols，amplificationprimers，PCR Ｐα/Ｍ，､〃α"αlbMS-MPtreeswerecon‐ protocolsandthoseofsequencingreactionsare structedbyPAUPRat（Sikes＆Lewis2001)， describedpreviouslyinArikawaandHiguchi ｗｈｉｃｈｉｓａｔｏｏｌｔｏｉｍｐｌｅｍｅｎｔｔｈｅＰａｒｓimony (1999)，ＤｅＬｕｎａｅｔａＬ（1999)，ＭａｅｄａｅｔａＬ Ratchetsearches(Nixonl999)withPAUP*,over (2000),andTsubotaandcoworkers（1999,2000, PAUP＊4.0blO(Swoffbrd2002)Jnanalysesby 2001a,ｂ)． PAUPRat,MPtreesweresearchedusingthePar－ simonyRatchetsearchstrategyintwenty200it- ＤａｊＭ"αｌ１ｙｓｅｓ erationruns Ｄａｒａｓａ－Ａｔｏｔａｌｏｆｌ８１７６cLgenesequences Mzx加川/MjAoodq"αｳﾉsな－MLtreeswere wereexamined(TableLwithDDBJ/EMBL/Gen- constructedwiththeNuｃＭＬｉｎＭＯＬＰＨＹｖｅｒ‐ Bankaccessionnumbers)．Thisdatasetcontains sion23b3package(Adachi＆Hasegawal996)． ninesequencesobtainedfiomourlaboratoryand FortheMLanalysesHKY85model(Hasegawa l72registeredsequencesafterBLAＳＴｓｅａｒｃｈｅｓ etaLl985)wasusedastheestimatemodeLML ｉｎｔｈｅＮＣＢＩｈｏｍｅｐａｇｅｆｂｒtheDNAdatabase・ treeswereconstructedwiththelocalrearrange- ThematrixincludeslOOspecies（l020TUs)of mentsearchfiomtheNJtreebyNucMLand theHypnales,S1speciesofLeucodontales,４spe- NJdist1.2.5(Adachi＆Hasegawal996);ａｎｄＭＰ ciesofHookeriales，andothersasoutgroups，to ｔｒｅｅｓｂｙＰＡＵＰＲａｔ・Best-confbnnedtransition／ makeasurveyfbrthephylogeneticrelationships transversion（Ts/ＴＶ）parameterswereestimated betweenthehypnobryaleanmosses．〃"伽α/lリノー basedoncalculationswiththeHKY85modelby ヅomeかｊｃａａｎｄＰ/iWo〃かeﾉﾉａｐａｒｅ"swereused NucMLemployingthe“-topt，，optionparameter、 asthemostdistantoutgroups． 乃ｅｅｃｏ"ZparjSo〃ＭﾉﾉMog-ﾉﾉﾉﾋﾋﾞﾉﾉ/ＩＯＯＣ/αﾉ?‘sev- 』/jg71me"ノーManualalignmentwascarriedout e7α〃esな一TreecomparisonwiththeMLcriteria fbrthel81sequencesagainstthoseofPﾉiﾘﾉSCO〃－ byNucMLwascarriedouttoevaluatetheresult- 〃e"αｐａｔｅ"ｓａｎｄ此"co6r〕/"碗ｓｃａ６/ｗ"zwithan ingtrees、Severalmeasureswereusedestimating editorprogramandouroriginalprogram，Sclean thelog-likelihoodofdifferenttreetopologies， 281(byTsubotal998-2002,inourhttpsite,see standarderrors(SES)ofthedifferenceinlog-like- AppendixB）Theotheroriginalprogramswrlt- lihood(Kishino＆Hasegawal989),andtheboot‐ teninPerlandCwerealsoappliedfbrthedata strapprobabilityestimatedbytheＲＥＬＬ(resam- processingUndeterminedsites，gaps，andre- plingofestimatedlog-likelihoodofsites)method gionsnotclearlyalignablefbrallsequenceswere （KishinoetaLl990;Hasegawa＆Kishinol994） excludedfi･omtheanalyses、Amongsequences TheprogrampackageCONSELO1e（Shimo- 652 HikobiaVoLl3,No.４，２００２ daira＆Hasegawa200Dwasalsousedtocalcu- Ｐ/iﾘﾉﾉoge"αｊｃａＭ〕入Fes latesthep-valuesoftheconfidencefbrthebifUr- Thedatamatrixwaspreparedfbranalysesof catingcandidatetopologiesusingseveraltesting phylogeneticrelationshipsofthehypnobryalean procedures：theApproximatelyUnbiased（AU） mosseSAtotalof2,223topologies，ofwhich test(Shimodaira2000,2002)usingthemultiscale only7treetopologieswerebifilrcate，ｗｅｒｅob- bootstraptechniquqtheKishino-Hasegawa(ＫＨ） tainedfTomfburanalyses：１ＭＬａｎｄｌＮＪｔｏｐｏｌｏ－ test（Kishino＆Hasegawal989)；andthe ｇｉｅｓｂｙＮｕｃＭＬ;ａｎｄ２，２２１ＭＰｂｙＰＡＵＰＲａｔｏｖｅｒ Shimodaira-Hasegawa（SH）test（Shimodaira＆ ＰＡＵＰ*・Onthe7bifUrcatingtopologies，local Hasegawal999;GoldmanetaL2000)． rearrangementsearcheswereconstructedfbrthe Ｂｏｏｍｍｐ卿ｊ－ＵｓｉｎｇＮｕｃＭＬ，abootstrap- obtainedtopologies・Atotalofl2bifhrcatingto- pingtestwithlocalbootstrapprobabilities(LBPs； pologieswereusedfbrthefbllowinganalysis、 iｎ％)(Adachi＆Hasegawal996)wasappliedfbr Moredetailedtopologiesweresearchedthrough ▼ thebestMLtopology・LBPisarelativebootstrap theobtainedtreesusinglog-likelihoodmeasure fi･equencyobtainedffomtopologysearchbylocal andseveraltests(Table２）HKY85modelwas rearrangementsofNucMＬＴｈｅｖａｌｕｅｗａｓｃｏｍ‐ usedastheestimatemodelwiththe 勺 parablewithavalueofFelsenstein1s(1985)boot- transition/transversionparameterestimatedけom strapprobability,anditisalittlelargerthanthe datasetfbrtheMLtreeas339・Thehighestlike- standardbootstrapprobability lihoodtreeofhypnobryaleanmosseswasob- Cb"Ｓｃ"s"ｓ／ree-Astrictconsensustreelbr tained,andshowninFigLTheLog-likelihｏｏｄ thebifUrcatingtopologieswithhigh-ranking ｖａｌｕｅｆｂｒｔｈｅＭＬｔｒｅｅｗａｓ－１５４４５８±794.6． log-likelihoodvaluesthatpassedtheAUtestwas ＴｈｅＬＢPsfbrthecladeswerecomparativelylow alsocoｍｐｕｔｅｄｂｙＰＡＵＰ*． values（41-100％）Becauseeachtopologydid notgainadecidedadvantageoverothertopolo- Results gies，ａ５０％majority-ruleconsensustreeof highrankingtreeswasalsoobtained，ａｎｄpre- Ｏ６ｊａｊ"ecl/se9"e"cedma sentedinFig2Thistreealsohasatendencyto- Newr6cLsequenceswereobtainedfbreleven wardtheMLtree・ species：ＢｊｓＭｊａノノ"g"〃α，ＣＭＯ"e"'o〃ﾉｶﾞﾉﾉcノー TheMLandconsensustreesfionnnotable """０，Ｇｏﾉﾉα"/α'wgj"Csα，Ｇ､Ｓ〃た"｡e"s,ｆ/ｏ"zα/ja- clades:(1)twomaiorcladesareshowninFigl cﾉellp〃sjaXgj"o"ｊα""８，〃ｂ"､ﾉﾉo〃dro〃scallpeﾉｰ [thecombinedHypnales（s・ｓｔｒｌ＝sensuVitt ﾉ(/bﾉﾉ川Ｌｅ"COC/ｂ〃Sc〃ｏｊ伽,ノVbcke'QpsjMjjj- l984D-Leucodontalesclade(87％LBPvaluein 伽ﾉﾋJ,Ｐｊ""areﾉﾉαα'、g"α、ﾉjα"z"o6'測加ｓ"6se‐ MLtree,８２％supportinconsensustree);andthe γj川"ＭｎｄＷ/iiﾉﾉImco"cav(/Ｍα，For〃ﾉhiaco" Hookerialesclade(64％LBP,７３％)]inthehyp‐ cav肋/jaanentiresequencel,428bplongwas nobryaleanclade(96％LBP,100％);ａｎｄ(2)４８ obtainedThesequenceshavebeensubmittedto basicclades[Sematophyllaceaeclade(96％LBP， theDNAdatabaseundertheserialaccession １００％)；Ｅ川血〃clade（96％LBP，100％)； numberｓａｓｓｈｏｗｎＴａｂｌｅＬ Ｅ"joc/､〃ノｗ６ｊｃ""c/"sclade；７１ﾉﾉ"城""Tclade (100％LBP，100％);肋ｊｅｔｊ"eﾉﾉａａ６ｊｅｊｊ"αclade； 比9"e"Ｃｅαﾉﾉg"ｍｅ"j AOpjerygj叩sjs-O"hotAecj"ｍｃｌａｄｅ（94％LBP， Thedatamatrixwaspreparedfbrphylogenetic 100％);ＥＣ"/ｑＷ〃"e"jjclade;Hypnaceaeclade analysesbymanualalignmentofthesequences・ (71％LBP，５５％)；血"COC/ｂ"-ne7o6r〕ﾉo''－０〕ノー Ｔｈｅｄａｔａｓｅｔｓｗｅｒｅｓｕｂｍｉｔｔｅｄｔｏａ５％chi-square p力aeaclade(89％LBP,５５％);Ａ"o"ZodO〃'wgeﾉﾉﾉ testbyTREBPUZZLE50(Strimmer＆Haeseler clade；Ｈｅ”ogje"αｐｅ〃o6"sjaclade；んOp/eﾉﾌﾉｰ l996),andallthedatapassedthetest・Forthe gj""ﾊﾉﾉ"eα/eclade;Prjo"odO〃(ノセ"s"sclade;。" analyses,atotalofLO92sitesofl8176cLse- /伽cﾊﾉα／b""Csα"αclade；Ｒﾉﾉ〕ﾉ肋α火llpA"s- quenceswereusedNucleotideｆｉ･equenciesesti- fMoco〃"m-LoGWo6/Ｗｍ－Ｃｔｅ"/血"mclade(８９ matedftomthedａｔａｓｅｔｗｅｒｅＡ＝29.2％，Ｃ＝ ％LBP,100％);ＭﾉＭ"碗-sb〃o〃"mclade(８８ 17.5％，Ｇ＝22.2％，ａｎｄＴ＝31.1％．Inthesites ％LBP,１００％);伽c/iDlp"s-P叩ﾉﾉﾉarjq-伽c/1)ノー ofthealignedr6cLsequences,４１７sites(＝３８２％ podopsjs-D"/ﾊﾉeﾉﾉａｃｌａｄｅ（89％LBP，100％)； inLO92sites)werevariable． P/cJgjotAecj""ｚｃｌａｄｅ（98％LBP，１００％)； Ｈ・TsuBoTA,Ｔ､ARIKAwA,Ｈ・AKIYAMA,ＥＤＥＬｕＮＡ,Ｄ・GoNzALEz,Ｍ・HIGucHIANDH・DEGucHI ６５３ Table2Compansonoflog-likelihoodscoresamongthel2resultingbifilrcatingtopologiesamongthe hypnobryaleanmossesfbr76cLsequenceswiththeHKY85model(HasegawaetaL1985）Thelog-likelihood valueswerecalculatedbyNucML(Adachi＆Hasegawal996),andallthep-valuesbyCONSEL(Shimodaira＆ Hasegawa2001）fiPomlO,O00repetitions、Thelog-likelihoodvaluesofthehighestlikelihoodtreesaregivenin anglebrackets,andthedifferencesinlog-likelihoodofalternativetreesfiomthatoftheMLtreeareshownwith theirstandarderrors(SE)fbllowing±・Thep-valuesthatarenotsignifiCantata＝OO5areemphasizedinbold typeTopologieswithasterisksintheAUcolumnwereusedfbrtheconsensustree(Ｆｉｇ２） Ｒａnｋ△/士ＳＥ MethodProgram Ｎｏｔｅ ＡＵＢＰＫＨＳＨ 。ｇ傘４白５●６竃３傘８●３戦０傘７△４。２３ 砂茄佃瓠⑬州Ⅳ列ｍ刈りⅡ ００００００００００００ ２００１０１００００００ ３８９１５４４４５７４０ ９０２５５６８５７８５０ ００００００００００００ ６３３３２３２２２２１０ ５４０５９４８７７５７３ ４６５８０８３７６３９０ ００００００００００００ ９８８８８７７７７６４０ ５９５４１７９７３０８４ ７５８６２９５４０１１６ 皿ⅢⅢⅢⅢⅢｗｗｗⅢⅢ川 ００００００００００００ 引叺２６６０２３孔０４０ ５０シ７５０シ０シ５（Ｕ１４００”５ 斜士士士士士士士士士士士 ５２２４３５３３４０６〆０８ ８６８Ⅱ０４９８Ⅱ。Ｌ０Ｏ ＲＲＲＲＲＲＲＲＲＲＬＬ 馳即岬山川川岼岼岬岼 醐蜘ＵＭ皿皿刀刃勺刃 ＬＬ 眺臥恥恥恥恥叺叺恥恥叩、 ｍｍｗｍｍｍｗｍｍｍ叩珊 ＋＋ｌ＋＋＋ｌＩくＩＮ シ７〈Ｕ今Ｊ７７６（５６４，句Ｊ７』 ｔｔｔｔｔｔｔｔｔｔ く－１１１２２つ』２４‐５６ ａａａａａａａａａａ ●●●●●●●●●●●● ＭＬ(Fig.1） ●●●●●●●。●●●● ●●●●●●●●●●●● LRSfiPomMP-l ２３４５６７８９０１２ LRSfiDmMP-2 ’’’’’’一｜｜刊 LRSfiPomMP-4 LRSfiFomMP-3 LRSfiPoｍＭＰ－５ ◆ ＭＥ １１１ ＬＲＳｆｉＰｏｍＮＪ－ｌ ｄ Ｊ 川 Ｊ ＋ Ｓ Ｎｏｔｅ－ＡＵ:Approximateunbiasedtest(Shimodaira2000,2002),ＢＰ:thebootstrapselectionprobabilityofFelsenstein（1985） estimatedbytheRELLresamplingmethod(KishinoetaLl990;Hasegawa＆Kishinol994),ＫＨ:Kishino-Hasegawatest(Kishino ＆Hasegawal989),ＳＨ:Shimodaira-Hasegawatest(Shimodaira＆Hasegawal999;GoldmanetaL2000);ａｎｄＬＲＳ:thelocal reanPangementsearch(Adachi＆Hasegawal996)ｂｙＮｕｃＭＬ P/e"rozj"ｍ-町/ＯＣＣ〃ａｓ伽"ｚｃｌａｄｅ（82％LBP， Oursmdycorroboratesthatl)theHypnales(s 82％);Brachytheciaceaeclade(66％LBP,１００％)； strl＝sensuVittl984]）andLeucodontalesare Climaciaceaeclade(99％LBP'100％);Ｐ/e"rozJ- notmonophyleticentities，whiletheHookeriales OpsLsclade;Stereophyllaceaeclade（１００％LBP， withtheinclusionofthePtychomniaceaeara2） 100％);lVboC/O/jc/io"ｚｊｊｒａＷ""α"e"sjlyclade;〃 thecombinedHypnales（ｓｓｔｒ．［＝sensuVitt ｍｅ"卯""ｍ〃"e"sclade；Amblystegiaceaeclade l984]）ａｎｄLeucodontalescompriseawellsup‐ (99％LBP,100％);Ｈｊｌｐ""、ｃ叩ressl/M71eclade； portedsistercladetotheHookeriales;ａｎｄ３)the 伽c/iﾘﾉﾉo加ａｊＭｃ"mclade；Neckeraceae-FoKMo- familiesEntodontaceae（s・str．［＝sensuVitt Ｐ e〃αclade(86％LBP,100％);Lembophyllaceae l984;TsubotaetaL2000]),Fontinalaceae,Ｓｅ‐ clade（82％LBP，100％)；nJxjp/iし〕ﾉﾉﾉ"m-G/Cs‐ matophyllaceae（s」at.［＝sensuTsubota＆co- Sα(/elp/i"s-Mi〕ﾉa6ea-肋Ｍｊａ－Ｈｏｍα/ｊａｄｅ/p伽 workers2000,2001a,ｂ]),Thuidiaceae(ｓｓｔｒ.[＝ clade（90％LBP,７３％);Anomodontaceaeclade sensuTsubotaetaL2000DandPlagiotheciaceae (70％LBP，１００％)；Fontinalaceaeclade（100％ (s・str.［＝sensupresent]）areresolvedasmono- LBP，100％)；Hookerialesclade（64％LBP， phyleticgroups、MembersoftheAmblystegia- 73％);LeY）/Cs/ｏｍ"mmacrocaZp"mclade;Ｂａ7〃－ ceae，HypnaceaaandLeucodontaceaearedis- 〃ａｐｏ加加'伽clade；Cyrtopodaceaeclade persedamongseveralclades． (100％LBP，100％)；HDlp"ode"Q/、〃〃?e"z/asi/ clade；RacOpj/""ｚｃｏｍ′ｏｍ/ace""Zclade；けr‐ Discussion r/io6M"、ﾉαﾉﾉjs-g7aZjaec/αcノビ,．』"/ZJco"?"j""?／z"‐ gjc/""zclade；Ｏ〃hoc/Ｄ"〃"１ノノ"eα'でclade； LOrdinalrelationships P/iﾉﾉo"otjs-Leｊｏｍｅ/αclade（97％LBP，100％)； Ourphylogeneticanalysiswith76cLlarge-scale Ｍｅﾉﾉcﾊﾉjq/ｂｒａｅ/o"gajeclade；PAO/ｊａｃｊｗ血 datasetdidnotrecovereitherthemonophylyof cladaMniaceaeclade（100％LBP,１００％);Hed‐ theorderHypnales(s・str.[＝sensuVittl984]),or wigiaceaeclade(100％LBP,１００％);ａｎｄＰＷｔＪｍｌｊａ themonophylyoftheLeucodontales；although ｈ〕lgromeかjcaclade］ theanalysisrecoveredthemonophylyofthe 654 ＨｉｋｏｂｉａＶｏｌｌ３,No.４，２００２ 姜霧iiil1ili篝liiilii蕊'１ ｐﾉapo､'cum 霧 蕊》 ７ _rｌ 鱈 １ 建霞一 一・・ 〕０ アートど１７５◆ ：．＆ｄご■訳 △●甲 ５７M鰯M雛'p"a"。'DaCea 田口 ＵＦＵＵＸＩ oｈａｍａｅ で ５７ 8７ iLL蕊llil1m ｣凸 ｎｄＩ:ＦＷｒ/世ＴＩ 鑿 Hypnales(slat） [Hypnales(ｓｓｔｒ.)‐ 潟溌iMIlmIMH LeucodoniaIes】 ■「 品;HMI鵲.ｌｉ８Ｉ;X： ユ品 ０ 空室竺彗i護篝iliIiiiiliii1iW鱸 ｐＨ ０.O1substitutions/sltE FiglThehighestlikelihoodtreefbrthealignedl,Ｏ９２ｂｐｏｆｔｈｅｌ８］ノカcLgenesequences(HKY85model;α/β＝ 339;ｌｎＬ＝-154458±794.6ｂｙＮｕｃＭＬ);andthemajorclades:theHypnales(s」at.)clade[＝thecombined Hypnales(s・str.[＝sensuVittl984])-Leucodontalescladc],andtheHookerialesclade・Ｔｈｅｈｏrizontallengthof eachbranchisproportionaltotheestimatednumberofnucleotidesubstitutions・Therootisarbitrarilyplacedonthe branchleadingtothenl"αｒｊａﾉl【Woll1eﾉﾉ化αａｎｄﾉ､ﾉﾘWol"/Ｍ/αﾉﾌqZe"s、Localbootstrapprobabilities（LBP;％） morethan50％areshownaboveornearbranchesShadedboxesidcntifythetraditionalLeucodontales． HTsuBoTA,Ｔ・ARIKAwA,Ｈ・ＡＫＩＹＡＭＡ,ＥＤＥＬｕＮＡ,Ｄ・GoNzALEz,ＭＨＩＧｕｃＨＩＡＮＤＨ・DEGucHI６５５ （・←、一・の）の①而亡・夛工 の①ｌｍＥ○つＯＮ一匹 の①而己囚 ‐（・』←の・の）の①一⑩亡旦夛エ］ 。… 認’三 園ＩＣ ク の②直のＥＯ ②の己－０亡巨。、 Ｂ 霊§甑： ←筐．。 ・こり二コ。：ごりこ３ヒー ｉ 藻霞'１蕊:： 篝聴 幻の亘旦Ｅ”Ｅ、。＆のＥＳ①ＪＩＩ５０ＩＩＩｌＩＩＩＩＩＩｌｌⅡＩＩＩＩ 言◎のＥｏミＥミ匡二０１１１１１ⅡＩＩＩｉ０ｌＩＬｇ← Ｕｏ』、」 ｏｏＥｍ ｍピヨのｌ ロ、 巴ｍｐＥＱＣｍ亡哩◎一望》記艶斗至禧陰曜哩 冨鷺蔓駕罵 ： ⑩巨岩Ｅ、の二ＯＥＣ壹窯込０ⅡⅡ 米 房一望 【壱○ llllililil 。Ｑ巴Ｅｍ橲而 ili の① 、 割ろ ｑＤ ＧＯ － ｇ M'塾!:!L1,,,蝿j雲璽！'wi塗：蝕幽鍵塾蝋！ MLllj 軸ロ IU ム 1１ illI …… FigLcontinued 656 HikobiaVoLl3,No.４，２００２ TUL ;:柵;脇|脇:脇 mj;!： ;i1fWl3:閉iWE;:鰐 TrIiiiE １０Ｃ 】【 ｅｃＩａｄｅ 虐門 鑿 ゴ ■■ ご『ロ １－ｺ Ｑ 』 津 繍灘 ｺﾞueWH認1N;。｡｡。 崖 Ｊ【Ｊ【Ｊ「Ｊ【二「【】信 Entodontaceae(ｓｓｔ｢.） 一一一 蝋ilj鑿iil識w二二＝． ThuﾉdjUmclade smuel/ｍａｎａ 梛liiﾄ|(;lmll;;』 ｍねscens-． ﾉsOpfelygjOsjS-O'tholhec/umclade 鑿蕊llii鑿二 Hypnaceaeclade 蝋i霧ｌＩＩｉ曇り率 繍iiiii1il1ji`。 LeucodOn-PleloblyoかＣ'yphaeacIade 蕊繍 け〃ごｌｓ ｉｊｓｏｂ "ａ ｄｂｎｔ ｂｕｓＣ， ，ｎＳＵＦ Ｉｏｎｅｌ subspyuennanensls ｇｅ〃 ｇｎＤｂ ＶＴｎｅ届 A､"ｍＣｈｊａｌｂ'mosana １００ ００Ｆ 5５ 2182 Rhyl/diade/Phus-ﾉヵﾉbcomjUm‐ ljillll讓蕊露i簔 LoeskeoblWm-Clenjdjumclade 65119° 」』且 Myu"um-Scjammjumclade １００ ００「 77achypus-Papjｿlalja- １００ ７７achypodOpsjS-Dulhje"aclade ，１，ii1； P/agjOfhec/umclade 嘩礎砿靜 鑿 鑿|篝iiiiili義！ P/euloz/um-hMOcom/as伽ｍｃｌａｄｅ 鑿霧護讓襄 BrachytheciaceaecIade C1imaciaceaecIade 鑿iiliiiiii雲霧 StereophyIlaceaeclade Ｆｉｇ２Ｔｈｅ５０％majority-ruleconsensustreefbrthellbifUrcatingtopologieswithhigh-rankinglog-likelihood valuesthatpassedtheAUtest、Therootisarbitrarilyplacedonthebranchleadingtothe〃"Ｍα/1Ｗｏｍｅｒ７ｊｃａａｎｄ Ｐ/1〕ﾊFCC〃Ｍ/〃α/e"s、 Ｈ・TsuBoTA,ＴＡＲＩＫＡｗＡ,Ｈ・AKIYAMA,ＥＤＥＬｕＮＡ,Ｄ・GoNzALEz,Ｍ・HIGucHIANDH・DEGucHI６５７ ＊ ５５ 塔惟蝉嚥舗塑舗篤一控 ides 11霧簔繍鍵X AmbIystegiaceaeclade 6４ 鍵鵜 Neckeraceae-FO'ssf71oemjaclade ’ ＵＩＢ「ｺ 8２ ﾌｅｍｌ．、■ ｡ｎｍO月月行kＢｍ 、｡"ｃｈｏｍｌｆｎｏＤ LembophyIlaceaecIade ;鰯liil菫iil;蕊 耐xﾉPhyﾉﾉUm-G/Ossadelphus-Myabea- 8２ B'Sse脂一HOma脂deゎhuscIade AnomodontaceaecIade 1００ Fontinalaceaeclade 5５ ＵＵＫＵ『０．回ＵｕＩＩＩＵ ｉ鰯jH轍!’ Hookerialesclade 5５ ii1鱗ill蝋…, MaclDmlmUmjncuMわ"ｕｍ ＭａｃｍｍｊｍＵｍｊｎｃｕｒＷｂﾉﾉﾘ、 霧 7３ Cyrtopodaceaeclade 1００ 鰯lljll:鰹H1鯏但竺昌ニニニニニニコＰ'Wbno施一LejOme/aclade 糒噸一顎 階鰐WI:|i;!；a｡lbngaね １００ Mniaceaeclade ，．.「 |j篝鍵鍵il1篝 HedwigiaceaecIade |蕊lii霧i篝！｝ Fig.２．continued , HookerialesThecombinedHypnalesandLeuco- somespeciesoftheBryales,Hookeriales,Leuco‐ ▲ dontales，however，compnseawell-supported dontalesandHypnaleSDeLunaetaL(2000)and sistercladetotheHookeriales・Ourresultsalso BucketaL(2000)revealedthenon-independence showedthemonophylyofallhypnobryalean ofthetraditionalordersHypnalesandLeucodon- mossesaspreviouslydiscoveredbyDeLunaand tales,andsuggestedthenecessitytobereconsid- coworkers(1999,2000),andBucketaL(2000)． eredasasinglemajorlineage Inthepresentstudywithalarge-scaledataset， HDlp"α伽α"`Le"ＣＯ血"〃/“ monophylyofeachorderHypnales(s・strI＝sen- Tsubotaetal．（1999）showedapreliminary suVittl984DorLeucodontaleswasnotrecov- phylogeneticrelationshipoftheHypnalesbased eredunderdifferenttreesearchprocedures・The on76cLdataset，especiallyrevealingthepoly- combinedHypnales（s・str.）andLeucodontales phylyoftheHypnaceae，ｗｉｔｈ３４０ＴＵｓｆｂｒｔｈｅ cladewassupportedwith８７％ＬＢＰｉｎｔｈｅＭＬ ａｎａｌｙｓｉｓ、ＤｅＬｕｎａｅｔａＬ（1999）suggestedthat ｔｒｅｅａｎｄ８２％supportintheconsensustree,root- monophylyofpleurocarpousmossesincluding edbythecladeoftheFontinalaceacThiscom- 658 ＨｉｋｏｂｉａＶｏＬｌ３,No.４，２００２ binedcladecon･espondstotheHypnales(s」at.） HDP"αceae asrevealedbyDeLunaetal.(2000)andnamed ThefiamiliesHypnaceae（withca40genera： astheHypniaebyBucketal.(2000)．Members Vittl984)areamongthemostdiversifiedtaxon- previouslytreatedastheLeucodontalesappeared omicgroupsinthepleurocarpousmosses、The asseveralcladesinthecombinedclade・Ourre- ftlmilialcircumscriptionwithintheHypnales，es- sultsofthepresentstudybasedonthe76cLgene peciallybetweentheHypnaceaeandotherfami- donotconcurwiththeclassificationandphylog- lies，suchastheSematophyllaceae,Plagiothecia- enyoftheorderHypnalesproposedbｙＶｉｔｔ ceae，andAmblystegiaceae，ｈａｓｌａｔelybecomea (1984),andBuckandVitt(1986）Theseresults subjectofspecialinterestandbeenrepeatedly alsosuggestthattheirsporophyticmorphological discussed(e9.,NishimuraetaL1984)． similarities，suchasdevelopmentofperistomal Ｉ、theobtainedMLandconsensustrees，the teethandexothecialcells,aremultipletransitions familyHypnaceae,aswellasitstypegenusHDlp- @ tosimilarsolutionstoepiphytismintheHypnales ""肌provedtobenon-monophyleticdistributing (slat.)． invariouspositionsinthetrees,regardlessofthe inclusionofD/α伽α火!〃/1α(inc.〃ｏｒ/ieM/α)and HDCﾉ(ｅｒｊａにs 〃/hiainthefamily，althoughtwospeciesof MonophylyoftheHookeriales［inc・ｔｈｅＰｔｙ‐ fbip"z川ａｎｄｔｈｅｇｅneraGoﾉﾉα"jαａｎｄ〃/αjsja chomniaceae,representedbyPltycAom"jo〃αcjc"－ fbnnasinglecladewith71％LBPsupportThis ，７e;sensuBucketaL2000;sensuBuck＆Goffi- fact，togetherwiththepresentanalysiss叩ple- net2000]wassupportedwith55％ＬＢＰｉｎｔｈｅ mentedbyadditionaltaxa,corroboratesthereport ＭＬｔｒｅｅａｎｄ７３％supportinconsensustree，be- ofTsubotaetal(1999)，suggestingthatthefiam‐ ingsistertothecombinedHypnales-Leucodon- ilyHypnaceae,aswellasthegenusjL卯"""',is talesinourphylogeneticanalyses;althoughsome polyphyleticalthoughthegenushasbeencare- oftheobtainedtreesshowedthatthePtychomni- filllyrevisedbymonographers(e9.,Ａ､ｄｏ1986, aceaeappearedasasistertothelargecladｅｉｎ‐ 1995)．Theresultsofthepresentstudybasｅｄｏｎ cludingothermemberoftheHookerialesandthe ｔｈｅ７６ｃＬｇｅｎｅｄｏｎｏｔｃｏｎｃｕｒｗｉｔｈthecircum- combinedHypnales-Leucodonta1es・Moredata scriptionoftheHypnaceaebyNishimuraetal． areinneedtoclarifythephylogeneticpositionof (1984)．Thisalsomeansthatthetaxonomiccon‐ thePtychomniaceae， fUsionintheHypnalesmightbeascribedtodiver- ＴｗomajorcladesoftheHypnales（s」at.［＝ sifyoftheHypnaceaecurrentlyunderstoodasa ｓｅｎｓｕＤｅＬｕｎａｅｔａＬ２０００；sensuBucketal largetaxonomicgroupcomprisingalargenumber 2000D:thecombinedHypnales(s・str.)andLeuc- ofspecies、 odontaleswiththeexceptionoftheCyrtopo‐ Nishimuraetal.（1984)placedCZJﾉﾉﾉe堰o"eﾉﾉZJ daceae,andHookerialesfbnnasinglelargeclade inthesubfIamilyHypnoideaeofthefhmilyHyp- withstrongsupport(96％LBPintheMLtree， naceae．Hedeniis（1990）transferredH)lp""籾 １００％supportinthcconsensustreeLasrepre‐ ﾉﾉ"肋eZgjjMittintothegenusCMjeZgo"eﾉﾉα sentedinpreviousworks(DeLuna＆coworkers (Amblystegiaceae)basedoncharactersincluding l999,2000;BucketaL2000)． capsulestructure，alarceUsandpseudoparaphyl- lia、Ａｎｄｏ（1995)，however，disagreedwith 2．Familialrelationships，especiallytheHyp- Hedeniis（1990）andretainedH)lp""碗//"cIl6ergjj naceaeandpossiblyrelatedfamilies inthegenuM卯""mTheobtainedtreeshows hthepresentinvestigation,somefiamilies,such themonophylyfbrtheH1lp""腕ノノ"cIlbergjj-CtJﾉｰ astheSematophyllaceae（s」at.)，Plagiothecia‐ ﾉje堰o"eﾉﾉαｃ"Spjc/、αcladesupportingbyhigh ceae（ｓｓｔｒ.）andFontinalaceae,areresolvedas bootstrapvalues（100％ＬＢＰｉｎＭＬｔｒｅｅ)，ａｌ‐ monophyleticgroups,whereasnoparticularclade thoughthecladechangeditspositionaccording isrecognizedtoaccommodatemembersofthe totopologiesThisresultwassuggestedby familiesHypnaceaeandLeucodontaceaeThe Hedeniis(1990)basedonthecharactersincluding Fontinalaceaeareplacedatthebasalpartofthe capsulestructure，alarceUsandpseudoparaphyl- combinedHypnalesandLeucodontalesclade． IiaandTsubotaetal.（1999)basedonlbcLdata； andconfinnedagaininthepresentstudy． HTsuBoTA,Ｔ・ARIKAwA,Ｈ・AKIYAMA,ＥＤＥＬｕＮＡ,Ｄ・GoNzALEz,MHIGucHIANDH､DEGucHI ６５９ 比marqp/1〕ﾉﾉﾉZJceae significantfbrtaxonomicreCognitionofthefam- TheSematophyllaceaeisalsoalargetaxon- ilySematophyllaceae,asconcludedbyTsubotaet omicgroupincludingnumerousspecies(withca aL(200ｌｂ)．Moreover,morerapidrateofnucle- 50genera:Vittl984),andthelimitsbetweenthe otidesubstitutionsoftheSematophyllaceaｅ（S， HypnaceaeandtheSematophyllaceaevaryamong lat.)wasdifferentfiomthatoftheotherlineage taxonomists，ＴａｎａｎｄＹｕ（1998,1999)madea and,therefbre,characteristictothislineage・This cladisticanalysisfbrthemorphologicaldata,and isolationmighthavebeencoITelatedtotheeco- showedtheparaphylyoftheSematophyllaceae， logicalrequirements，asthesetaxainthelineage Tsubotaandcoworkers（1999,2000,2001a,ｂ） withhabitatpreferencefbrgrowingontree clarifiedtheirphylogeneticrelationshipsinthe trunks． hypnobryaleanmosses，andshowedthemono- Ｐ phylyoftheSematophyllaceae（s」at.；sensu Ｅ'zroqlomaceae TsubotaetaL2000，2001a,ｂ)．Arikawaand Severaldiscussionshavebeenmadeonthere- Higuchi（2002),however,showedtheinclusion lationshipoftheEntodontaceaeandThuidiaceae oftheEntodontaceaewithinthecladeoftheSe- totheHypnaceaeandtheSematophyllaceaefiom matophyllaceae(slat.),althoughthesupportwas molecularviewpoints・Mizushima（1960）ａｎｄ notsostrong Buck（1980）publishedmonographicworkson DistinctionbetweentheHypnaceａｅａｎｄＳｅ‐ theEntodontaceaaandtheyhavedifferentviews matophyllaceaehasbeencontroversiaLespecially withregardtothesystematicpositiｏｎｏｆｔｈｅｇｅ－ concerningthephylogeneticpositionofsome ｎｕｓＯ７ｒｈｏ/hecj"腕．Mizushima（1960)placedit genera,suchas〃/α〃α火ＩＩＰＡα(incB7oj/je7eﾉﾉα)， inEntodontaceae,whileＢｕｃｋ(1980)intheHyp- 〃伽ａｎｄＨｂｊｅｍｐＭ/i"腕．ＤｅＬｕｎａｅｔａｌ． naceae，Ｖｉｔｔ（1984)，andBuckandGofTmet (2000)ａｌｓｏshowedtheroughconceptoftheSe‐ (2000)treatedthefamilyinanarrowsense,in‐ matophyllaceae，althoughAQprer〕/ｇｊ"ｍｒｅ"e'wm cludingonlythefburgenera:Ｅ"加巾"，ErWAro- wouldneedtobetransfelTedfiomtheHypnaceae、 c/o"〃腕，MCso"oc/O〃ａｎｄ〃/伽job'Ｗｍ．The ArikawaandHiguchi（1999）suggestedD/伽ja presentsmdyshowedthattheEntodontaceae（s polb′α"伽reported，ａｓ〃/α伽eﾉﾉａｐｏﾉ!〕ﾉα"伽， str.)confinnstheexclusionofO7tAo/hecj"ｍａｎｄ shouldbeconsideredintheSematophyllaceae ＲＤｅ"血Sc/empocﾉﾉ"ｍｆｆｏｍｔｈｅｆａｍｉｌｙ,supporting OurresultshowedthemonophylyoftheSemato- thetreatmentofVitt(1984),andBuckandGoffi- phyllaceae(slat.)proposedbyTsubotaandco- net（2000）ｏｒ/hoZhecj"腕ノW/bsce"sfbrmeda workers（2000,2001a,ｂ),includingthegenera cladewithhOprerygmp川加"eﾙｧﾉα"α,and D伽jα火ＩＩＰﾉｶα，附加ａｎｄＨｅ〃qp/Ｍｊ川and 囚ye"dmc/empodj"ｍｐ"7"mappearedintheclade somespeciespreviouslytreatedasmembersof withmembersoftheBrachytheciaceae、Thetree theHypnaceae：Ｈｊｌｐ""加伽ro-v"枕,ZFOpjer〕ノー alsoshowsthatthecladeco､sistingoftheSe- ｇｊ"腕／e"e'ｗｍａｎｄ〃/αｊｓｊａｐｏﾉ!〕ﾉα"ﾉﾉ､,ｗａｓｒｅ－ matophyllaceae（s」at．［＝sensuTsubotaetaL ｖｅａｌｅｄｂｙｂｏｔｈｔｈｅｍｏｓｔＭＬｔreewithstrongsup- 200q2001a,ｂ]),EntodontaceaeandThuidiaceae port(96％LBP)andtheconsensustreeWecan- wassupportedwithhighLBP(97％LBPinML Ｐ notsupportthephylogeneticpositionofP､pobノー tree)．Vitt(1984)placedtheEntodontaceaeclose α"rAaonthebasisofthedatapresentlyavailable totheSematophyllaceaeonthebasisofmorphol- Further,itisnecessarytoverifytheidentification ogicalandecologicalcharactersOurresultsup- ofthematerialofthespecies・ portsthetreatmentoftheEntodontaceae,butnot Aspreviouslyindicatedinmorphologicaldata theThuidiaceae． analysesbyHedeniis（1995,1996),phylogenetic relationshipsbasedonmoleculardatashouldbe Ｐﾉagjoj/iecmceae surveyedbetweentheSematophyllaceaeandthe Thetreeshowedthecladeconsistingofonly familiesintheorderHookeriales・Themono- thegenusP/αgjorhecj"mnearthecladesinclud- phylyoftheSematophyllaceaeandtheHookeri- ingtheBrachytheciaceaeandsomemembersof aleshadnophylogenicevidenceinouranalyses theHylocomiaceae・Recently，Pedersenand Thepresentsmdyrevealsatleastthatthechar- Hedeniis(2002)showedthephylogenyofthePla- actersofwell-developedalarcellsandhabitatare giotheciaceaebasedonmorphologicaｌｄａｔａａｎｄ 660 ＨｉｋｏｂｉａＶｏＬｌ３,No.４，２００２ twoDNAregions:ノアｓ４ａｎｄ伽Ｌ－伽F,suggesting ogyandtreatedtheAmblystegiaceaewithanar- thefamilywithawidecircumscriptionArikawa rowersenseThepresenttreeconfinnsthisnar- andHiguchi(2002)clarifiedthephylogeneticpo‐ rowefamilialcircumscription，placing4"αcam- sitionofthegeneraA0prerygj叩巾andHblzo- ptoqlO〃intheAmblystegiaceae． gjeﾉﾉα，andproposedtheinclusionofAOpZer)Igj- qpsなwithinthePlagiotheciaceaeThepresent 〃αc/iﾘﾉrAecjaceae investigationdidnotsupportthesepreviousanal- Thepresentsmdyshowedthemonophylyof yses,discoveringthemonophylyofthePlagiothe- theBrachytheciaceae，includingOAzJ〃"ｍ１ｅａａｎｄ ciaceaeincludingonlythegenusP/αgjorAecj"腕． Ｈｂ/jco血""z"'ｚａｓｗｅｌｌａｓＲｓｅ"ﾛﾉＤＳＣ/empoい""Z andexcluding7bme"卯""腕.Ｈセノjcoc/Wz伽mwas S〃e叩Ｍ/αceae previouslytreatedintheMyriniaceaebyVitt ArikawaandHiguchi（1999),andthefbllow‐ (1984)andBuckandGoffinet(2000)．Ourresult ingstudies（e9.,Pedersen＆Hedeniis2002； supportstheplacementoｆＡｅ"cﾉＤＳＣ/empo`j"m Arikawa＆Higuchi2002)clarifiedtheexclusion byBuckandGoffinet(2000),ｂｕｔｎｏｔＨｂ"ＣＯＣ/b"‐ oftheStereophyllaceaeffomthePlagiotheci- 〃'７０．ＴｈｅｇｅｎｅｒａＰ/αgjor/iecj"腕,Ｐ/e"rozj"mand aceae,andthemonophylyoftheStereophyllaceae fMoco〃ａｓﾉﾉwmwereplacedinacommonclade basedonmolecularphylogeneticanalysesThe withthetaxaoftheBrachytheciaceae(excluded presentresultcorroboratedthepreviousworks Tb"Te"/ﾉ!〕lp""〃〃"e"s）ｗｉｔｈ６６％ＬＢＰｉｎＭＬ showingtheStereophyllaceａｅａｓａｓｉｓｔｅｒｔｏｔｈｅ ｔｒｅｅ． ７ｂｍｅ"卯""加一 ﾉVbo巾ﾉﾉc/ＩＣ加加clade，althoughtheirphyloge- MCjeo7mceae neticpositionisnotsostableintheHypnales(s、 HuttunenandIgnatov(perscomm.)suggested lat.[＝sensuDeLunaetaL2000;sensuBucket thecloserelationshipbetweentheBrachythecia- aL2000])clade． ceaeandMeteoriaceaeintheirpreliminalyanaly- ses・Ourpresentanalysisincludesonlyonespe- HMocomaceae ciesoftheMeteoriaceae・Furthersequencesof lntheMLtree,theHylocomiaceaeappearedas theMeteoriaceaeareneededtoshowtherelation- twodistinctclades:Ｒ/Uﾉ肋α火llpAMD-fMoco加川 shipsbetweenthetwofamilies． 一LoalAeo6'Ｗｍ－ｄｅ"jcﾉﾉ"腕cladeatthebasalpo- sitionofthecombinedseveralclades,suchasthe 、"伽aceαｅα"ＣＭ"o"zodo"raceae BrachytheciaceaecladeａｎｄｔｈｅＰ/CJgjorAecj"m TheThuidiaceae(s・str.[＝sensuTsubotaetaL clade;ａｎｄＰ/e"'℃ｚｊ"ｍ－Ｈｙ/ＯＣＣ〃ａｓ〃ｍｃｌａｄｅａｓ 2000]),consistingofonlythegenusT1h"ｊ`ｊ"川， ｔｈｅｓｉstercladetothePmgjoj/iecj"mcladcAre- fbnnedasinglecladebeingsistertotheEntodon- viewofthcirfamilialcircumscriptionisneeded taceae(s・str.[＝sensuVittl984;sensuBuck＆ １ fbrthegeneraP/e"７ozJ"mandfMoco〃ａｓ加川． Goffinet2000])，ａｓｓｈｏｗｎｂｙＴｓｕｂｏｔａｅｔａＬ (2001a)．肋ｊｅ""eﾉﾉZza6jerj"αａｎｄＢｏ"/ZlO′α〃ﾉｰ 伽6(Wegjaceae /emappearedoutofbutcloseto,thecladeofthe Thephylogenetictreesinthepresentstudydis- Thuidiaceac coveredpolyphylyoftheAmblystegiaceae（s Thegenera比p/o/j〕ﾉ腕e"伽ａｎｄ』"omo巾"， lat.）aspreviouslysuggestedbytraditionaltax- ｅｘｃｅｐｔｆｂｒＡｇｉ７ａｒＺﾉﾉﾉ，appearedinthecladein- onomists（e､９，Hedeniisl997)，showingthree cludingmembersoftheNeckeraceae，fbrminga clades:ＣＭＯ"e"ro"-DreYﾌα"oc/αc/"s-Hyg7oα"'‐ resolvedcladeplacingthesecondalybasalposi- 611〕Mgj"〃一A"αcα叩/o血〃clade,ＣａﾉﾉjeZgo"eﾉﾉa tioninthecombinedHypnales-Leucodontales clade,ａｎｄ７１ｏｍｅ"卯"""Zclade，BuckandGoffi‐ MaedaetａＬ（2000）alreadyclarifiedthepoly‐ net(2000)treatedthefiamilywithanarrowSense， phylyoftheAnomodontaceae，showingaclade excludingsomegeneratreatedinthepresent withHZZp/o/i)ﾉｍｅ"j"耐,比Ｚｐａｊ"e"ro〃ａｎｄｓｏｍｅ analysis,ｅ,9.,CMjeHgo"eﾉﾉα(inc.ｆ卯""腕ﾉﾉ"‘/L Ａ"o"zo吻刀species・Thesefactsindicatedthat 6e卿/）ａｎｄｎｍｅ"卯""腕，ｆｉ･omthefiamily・ somegenera,suchasHZJpM〕ﾉ腕e"伽ａｎＭ"o- Vanderpoortenetal.（2002）showedaphyloge- mo血〃whicharesometimestreatedwithinthe netictreebasedonDNAsequencesandmolphol- Thuidiaceae,shouldbedividedintotwofamilies： HTsuBoTA,Ｔ・ARIKAwA,Ｈ・ＡＫⅣAMA,ＥＤＥＬｕＮＡ,Ｄ､GoNzALEz,MHIGucHIANDH・DEGucHI６６１ theThuidiaceaeandAnomodontaceae，support- tionoftheFontinalaceae． ingthesystemofMaedaeｔａＬ(2000),andBuck andGoffinet(2000)． qWQpodnceae TheCyrtopodaceaeweretreatedintheLeuco‐ 比"ＣＯ伽"mceae，CrXpAaeaceae，α"‘Pjero6r〕ﾉa- dontalesbyVitt（1984)，ａｎｄＢｕｃｋａｎｄＶｉｔｔ Ceae (1986)Jnthepresentinvestigation,theCyrtopo- MaedaetaL(2000)alsodiscussedthefamilial daceae,representedbyB“Ｃｈｅ'eﾉﾉﾉａ６７ｅｖｉ/bﾉﾉα,Ｂ、 affinityofFolTsrroe〃α,showingrelationshipbe- e/egzJ"伽加α,ａｎｄＣＷｍｐ"sMos"s,appearedin tweentheCryphaeaceaeandLeucodontaceae・As apositionoutsidetheHypnales(s」at.［＝sensu theytreatedonlytheleucodontaleanmossesfbr DeLunaetaL2000;BucketaL2000])-Hookeri- theiranalysis,thephylogeneticpositionfbrmem- alesclade,fionninganotableclade(100％ＬＢＰin ’ bersoftheorderinthepleurocarpswasnotso theMLtree）sistertotheHypnodendraceae・ clear・hthepresenttreesbasedonthelarge-scale BuckandGofY1net（2000）treatedtheCyrtopo- dataset，thetraditionalleucodontaleanmosses daceaeintheRhizogoniales,andthepresenttree fbrmfiourma]ｏｒａｎｄｓｏｍｅｍｉｎｏｒｃｌａｄｅｓｉｎｔｈｅ supportstheirplacementofthefamily． ｃｏｍｂｉnedHypnales-Leucodontales，supporting thethreecladespresentedbyMaedaetaL(2000)． ﾉVCcAe'αceaeα'１．ｋ肋QpﾉＭ/αceae ThemembersoftheLeucodontaceae，exceptfbr Theotherrnembersofthetraditionalleucodon‐ Fo'Fsj7oe〃α，CryphaeaceaeandPterobryaceae taleanmosses,previouslytreatedastheNeckera- fbrmacladewithinthecladeoftheHypnales(s・ ceae,appearedasamajorcladewith86％LBP lat.[＝sensuDeLunaetaL2000;sensuBucket support,asasistertotheLembophyllaceaeclade・ aL2000])．Fol《Ｗｏｅｍｚａ,amemberofthetradi- Intheclade，ＥｃＡｊ"o血"腕〃m6ros"ｍａｓｗｅｌｌａｓ tionalLeucodontales，isplacedinthecladenear Ａ"o"TOC/O〃ｇ〃αﾉﾋI/ｊｊａｎｄＦｏ応sｊ'Ce"zjaspp、were theNeckeraceaeTheseresultsdidnotsupport alsoincluded theappearanceoftheleucodontaleanfamiliesin thecombinedHypnales（s・ｓｔｒｌ＝sensuVitt C/〃αcjaceaeα"c/P/e"rozmpMﾉﾋJceae l984])andLeucodontalesbyBucketaL(2000)． TheClimaciaceaeandPleuroziopsidaceae fbnnedaweakclade(57％LBP,９１％)nearthe Ｆａ６７ｏ"mceae Stereophyllaceaelreland（1968）segregated TheFabroniaceaefbrmnonotablecladeinthe P/e"γozjqpsLsfiomtheClimaciaceaeintothemon- MLandconsensustrees，asshowntbeabove otypicfamilyPleuroziopsidaceaeNorrisandlgna- Furtherinvestigationswithadditionaltaxafi･om tov(2000)observedthestemsurfaceanatomyin othergenera，ｓｕｃｈａｓＦｃＪ６ｒｏ"ｊα，ＳｂＡｗ伽cﾊﾙeα， C/〃αcj"mandP/e"mzmp川,andsuggestedthat , ａｎｄＳｂｈｗｅなcAAcOpsjSareneeded． P/e"'ozmpsjSshouldbereturnedtotheClimaci- aceacOurtreesdonotconfIictwiththeirsug- Ｆｏ"""α/αceae gestiononP/e"rozjOpsjS，althoughthevalueof AlthoughonlyFo"/ｊ"α/jJspecieswerein- Ｐ thesupportisｎｏｔｓｏｈｉｇｈ cludedintheanalysis，theFontinalaceaeappear asasingleclade,atthemostbasalpositioninthe 3.Singlecostaanddoublecosta Hypnales-Leucodontalesclade，supportingthe Vitt1ssystem（1984）fbrtheclassiflcationof conclusionofVitt（1984）ａｎｄＢｕｃｋ＆Ｖｉｔｔ hypnaleanmossessuggeststhatthegroupswitha (1986）Ｂｕｃｋ＆Vitt（1986）constructedanew singlecosta，asrepresentedbytheBrachythecia- classificationfbrpleurocaIpousmosses，recir- ceaeandAmblystegiaceae，wouldhavediverged cumscribingtheHypnales（s・str.）includingthe fi･omthehypnaleanancestoratanearlyevolu- HypnodendrineaeandFontinalineaeOuranaly- tionarystage,fiollowedbythegroupswithadou‐ sessupporttheinclusionoftheFontinalineae blecosta，ａｓrepresentedbytheHypnaceae，Se- withintheHypnales，butnottheinclusionofthe matophyllaceaeandEntodontaceae,whichhere- Hypnodendrineae、Furtheranalysisincluding gardedasthemostrecentevolutionarystageof somespeciesoftheFontinalaceae,especiallyａｎ‐ theorderHypnales（ｓｓｔｒ.［＝sensuVittl984]） othergenusDjchel【Wlm,wouldclarifytheresolu‐ (seealsoTsubotaetaLl999;BucketaL2000)． 662 ＨｉｋｏｂｉａＶｏｌｌ３,No.４，２００２ lnthepresentinvestigation,taxawithadouble Englishsummary） costaappearinvariouscladesofthetree・This Arikawa,Ｔ､＆Higuchi,Ｍ１999．Phylogeneticanaly- resultsuggeststhatasinglecostaisanancestral sisofthePlagiotheciaceae(MuscOanditsrelatives character,andthatthereweremultipletransitions basedon/御6cLgenesequencesCryptogamie,Ｂｒｙ－ toadoublecostaTherefbre，ourresultsofthe ｏＬ２０:２３１－２４５． presentstudydonotconcurwiththesuggestion ＆Z002Phylogeneticpositionofthe proposeｄｂｙＶｉｔｔ（1984)，ａｎｄＢｕｃｋａｎｄＶｉｔｔ generaLsOpte'f〕/gjOpsjFand比ﾉazOgjeﾉﾉα（Musci） (1986)．OuranalysesconfirmtheHndingsbyDe basedonlbcLgenesequences・BryoLRes．（in LunaetaL(2000)andBucketaL(2000)thatthe press） Hypnales，togetherwithmostoftheleucodon- Buck,Ｗ,Ｒ」９８０AgenericrevisionoftheEntodon- taleangroups，fbnnamonophyleticsistergroup taceae．』・HattoriBotLab48:７１－１５９． tothehookerialeanlineage・Resultsalsoshow ＆Goffmet,Ｂ、2000．Morphologyandclas‐ ､ thatthefamilialcircumscriptionwithintheHyp- sificationofmosses・Ｉｎ:Ａ､Ｊ・Ｓｈａｗ＆BGofHnet nalesandLeucodontalesshouldbereconsidered． (eds.),BryophyteBiology,ｐｐ７１－１２３・Cambridge UniversityPress,Cambridge． －＆Vitt,，.Ｈ」９８６SuggestionsfOranewfa- Acknowledgment milialclassificationofpleurocarpousmossesTax- on35:２１－６０ Thispaperisthecombinedresultofaninterna- tionalcollaborativeeffbrt、Ｗｅｗｏｕｌｄｌｉｋｅｔｏａｃ－ －，Goffinet，Ｂ＆Ｓｈａｗ，Ａ・I2000a・Testing ｋｎｏｗｌｅｄｇｅＲ．Ｄ・SeppeltfbrcheckingtheEng- morphologicalconceptsofordersofpleurocarpous lishtextandhelpfnlsuggestions;BGoffinetand mosses(Bryophyta)usingphylogeneticreconstruc- W．Ｒ・BuckfbrredressingtheEnglishandfior tionsbasedon/r"L-ﾉﾉ侭"Ｆａｎ。'ips4sequences・ＭｏＬ ＰｈｙＬＥｖｏＬｌ６：１８０－１９８． theircommentsandsuggestionｓｏｎａｎｅａｒｌｉｅｒｖｅr- ＆Z000hNovelrelation‐ sionofthispaper;ＭＳ・IgnatovandSHutmnen shipsinpleurocarpousmossesasrevealedbycp- fbrtheircommentsandsuggestionsonthephylo‐ DNAsequences・BryologistlO3：774-789． genetictrees;andBEstebanezfbrhergreathelp ＣＯＸ,ＣＪ.,Goffinet,Ｂ,Newton,Ａ､Ｅ､，Ｓｈａｗ,ＡＪ＆ WeａｌｓｏｔｂａｎｋＳＭａｅｄａａｎｄＫ・ＫＣsugewith Hedderson，Ｔ，Ａ､12000．Phylogeneticrelation- whomwehavecollaborated，MHasebeand，T shipsamongthediplolepideous-altematemosses Nishiyamafbrtechniqueofexperimentandad- （Bryidae）inferredfiPomnuclearandchloroplast viceaboutphylogeneticanalysis,ａｎｄＮ・Minaka DNAsequences．BryologistlO3:224-241． andMMiyafbrusageofPAUPRat・Researchon ＤｅＬｕｎａ,Ｅ,Newton,Ａ､Ｅ，Withey,Ａ､，Gonzalez,，． molecularsystematicsofmosseswasfUndedby ＆Mishler，ＢＤ、１９９９．Thetransitiontopleuro- thefbllowinggrants:KAKENHl（Japan）grants carpy：Aphylogeneticanalysisofthemaindiplo- O9839031toH､Akiyama,Ｏ９０４１１６５ｔｏＪＭｕｒａｔａ， 、 lepidouslineagesbasedonr6cLsequencesandmor- ｌｌ６４０６９８ｔｏＨ・Deguchi，l3640707toMHigu- phology・BryologistlO2:634-650. chi，ａｎｄｌ３７４０４９０ｔｏＨ・Tsubota；ａｎｄＣＯＮＡ－ ,Buck,ＷＲ.,Akiyama,Ｈ,Arikawa,Ｔ､,Tsubo- ＣＹＴ(M6xico)grant27400-NtoEDeLunaet ｔａ,Ｈ､，Gonzalez,，.,Newton,Ａ､Ｅ＆Ｓｈａｗ,Ａ､Ｊ、 aL 2000．Ordinalphylogenywithintbehypnobryalean pleurocarpousmossesinferredfTomcladisticanaly- Literaturecited sesofthreechloroplastDNAsequencedatasets： ／ﾉ･"Ｌ－Ｆ,ノアｓ４ａｎｄﾉ･6cLBryologistlO3:242-256． Adachi,』．＆Hasegawa,M1996ComputerScience Felsenstein,Ｊ・l98LEvolutionarytreesfromDNAse‐ Monographs，Ｚ８ＭＯＬＰＨＹｖｅｒｓｉｏｎ２３・Programs quences：amaximumlikelihoodapproachJ、ＭｏＬ fbrmolecularphylogeneticsbasedonmaximum ＥｖｏＬ１７:368-376. likelihoodl50pp、InstimteofStatisticalMathe‐ 1985．Confidencelimitsonphylogenies：an matics,Tokyo approachusingthebootstrapEvolution39： Ando,Ｈ1986．SmdiesonthegenusHyp"""ZHedw． 783-791． （1V)Hikobia9:467-484 Fitch,Ｗ・M197LTowarddefiningthecourseofevo- 1995．ThegenusHip"""？(Musci)inJapanlL lution：Minimumchangefbraspeciflctreetopol- Nat・EnvirSciRes､８:67-99.（ｉｎJapanese,ｗｉｔｈ Ｈ・TsuBoTA,Ｔ・ARIKAwA,Ｈ・AKlYAMA,ＥＤＥＬｕＮＡ,Ｄ・OoNzALEz,ＭＨＩＧｕｃＨｌＡＮＤＨ・DEGucHI ６６３ ogy・Syst・ＺｏｏＬ２０:406-416． ｒ６ｃＬＭｏｌＰｈｙＬＥｖｏＬ７：377-393． Goffmet,Ｂ，Bayer,ＢＪ.＆Vitt,ＤＨｌ９９８・Circum‐ Maeda，Ｓ，Kosuge，Ｋ，Gonzalez，，.，ＤｅＬｕｎａ，Ｅ＆ scriptionandphylogenyoftheOrthotrichales(Bry- Akiyama，Ｈ、2000．Molecularphylogenyofthe opsida)inferredfiPomr6cLsequenceanalyｓｅｓＡｍ suborderLeucodontineae（Musci；Leucodontales） ＪＢｏｔ８５：１３２４－１３３７． inferredfiPom'6cLsequencedata・JPlantRes、 Goldman,Ｎ､,Anderson,Ｊ､Ｐ.＆Rodrigo,Ａ・Ｇ・Z000 １１３：Z9-38 Likelihood-basedtestsoftopologiesinphylogenet- Magombo，Ｚ・ＬＫ・ZOO2Thephylogenyofbasal ics、SystBioL49:652-670． peristomatemosses：evidencefiomcpDNA，and Hasegawa,Ｍ＆Kishino,Ｈ・l994Accuraciesofthe implicationfbrperistomeevolutionSystBot.(in simplemethodsfbrestimatingthebootstrapprob- press)． abilityofamaximumlikelihoodtree、ＭｏｌＢｉｏＬ Manhart，Ｊ・Rl994Phylogeneticanalysisofgreen , ＥｖｏＬ１１：1４２－１４５． plantノ"bcLsequences・ＭｏＬＢｉｏｌＥｖｏ１．３：’１４－ ＆Yano，Ｔｌ985．Datingofthe １２７． buman-apesplittingbyamolecularclockofmito- Mizushima,Ｕ」960．JapaneseEntodontaceaeJ・Hat- chondrialDNA．』・ＭｏｌＥｖｏＬ２２：１６０－１７４． toriBot､Lab２２:９１－l58 Hedeniis,Ｌｌ９９０（1992)．Taxonomicandnomencla‐ Murray,ＭＧ.＆Thompson,ＷＦ・’980．Rapidisola- mralnotesonthegeneraCZJﾉﾉﾉeﾉ1go"e"αａｎｄＢ'で‐ tionofhighmoleculａｒｗｅｉｇｈｔｐｌａｎｔＤＮＡ・Nuc MerjqLindbergial6：１６１－１６８． AcidsRes」９:4321-4325． -1995．Highertaxonomiclevelrelationships Nishimura，Ｎ､，Higuchi，Ｍ，Seki，Ｔ＆Ando，Ｈ， amongdiplolepidouspleurocalpousmosses-acla- １９８４Delimitationandsubdivisionofthemoss disticoverview．』・ＢｒｙｏＬｌ８:７２３－７８１． ftlmilyHypnaceaeJ、HattoriBot・Lab、５５:２２７－ 1996（1997）Acladisticevaluationofrela- ２３４ tionshipsbetweentheHookeriales,theSematophyl- Nixon，Ｋ・Cl999TheParsimonyRatchet，anew laceaeandsomeothertaxaLindbergia21：４９－８２ methodfbrrapidparsimonyanalysisCladistics 1997.Ａ、evaluationofphylogeneticrela- l5:４０７－４１４． tionshipsamongtheThuidiaceae,theAmblystegia- Norris,，.Ｈ､＆Ignatov,MS20000bservationson ceae,andthetemperatemembersoftheHypnaceac stemsurfacｅａｎａｔｏｍｙｉｎＣﾉﾉ"1ａｃｊＬｌｍａｎｄＰ/e"'ozJ‐ Lindbergia22：ｌＯｌ－ｌ３３・ 叩叩(Climaciaceae:Musci）Arctoa9:l51-154 Ireland,Ｒ､Ｒ・l968Pleuroziopsidaceae,anewfamily Pedersen，Ｎ、＆Hedeniis，L2002Phylogenyofthe ofmosses．』・HattoriBotLab31：59-64. 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Anopenreadingframe(ycfll)isevolutionarycoL ＆１９９９Multiplecomparisonsof servedfiPomRTCyanobacteriatotheplastidDNAs log-likelihoodswithapplicationstophylogenetic ofArchegoniatesandGymnospermsiｓＲＴｍｏｄｉ－ inference、ＭｏＬＢｉｏＬＥｖｏＬ１６：ｌｌｌ４－ｌｌｌ６ ＨｅｄｉｎｔｈｅｐｌａｓｔｉｄＤＮＡｓｏｆＤｉｃｏｔｓａｎｄｉｓｎｏｔｐｌａｓ‐ Ｓｈｉｍｏｄａｉｒａ,Ｈ､＆Hasegawa,Ｍ・Ｚ００１ＣＯＮＳＥＬｆｂｒ tomeencodedinRTMonocots．』・PlantPhysioL ａｓｓｅｓｓｉｎｇtheconfldenceofphylogenetictreeselec- l46:Z58-262 tionBioinfbrmaticsl7：l246-l24Z Lewis，ＬＡ.，Mishler，ＢＤ．＆Vilgalys，Ｒ、１９９７． Sikes,Ｄ・Ｓ＆Ｌeｗis，Ｐ､０．２００１．ＰＡＵＰＲａｔ:ＰＡＵＰ＊ Phylogeneticrelationshipsoftheliverworts（He- ｉｍｐlementationoftheparsimonyratchet,betasofi- paticae),abasalembIyophytelineage,inferredffom ware,versionLDepartmentofEcologyandEvo- nucleotidesequencedataofthechloroplastgene lutionaryBiology，UniversityofConnecticut， 664 ＨｉｋｏｂｉａＶｏＬｌ３,No.４，２００２ StolTs accessionnumber，localityandspecimennumber，or Strinmner,Ｋ＆Haeseler,Avonl996Quartetpuz‐ referencesource・LeskeaceaeD"ｊＡｊｅ"α叩ecjosLDsj"za zlingaquartetemaximum-likelihoodmethodfbr BrothexCardot，ＡＢＯ19467,Japan，Shikoku，Kochi reconstructingtreetopologies・ＭｏｌＢｉｏＬＥｖｏＬｌ３： Pref，Monobe-mura，Befilkyo，９５０ｍａｌｔ.，Ｊｕｌｙ27, 964-969． 1998,HAkiyamal4356(HYo),thissmdy;Amblyste- Swoffiord,，.Ｌ2002．ＰＡＵＰ*・PhylogeneticAnalysis giaceaeOmo"e"ﾉﾊﾟo〃〃c伽'’１(Hedw）Spruce，ＡＢ UsingParsimony(*andOtherMethods).Version4 Ｏ95270,Japan，Honshu,Yamaguchi-ken，Shuhou-cho， SinauerAssociates,Sunderland,Massachusetts Akiyoshi-dai，Ｃａ、ｌＯＯｍａｌｔ.，August２７，１９９８，Ｈ、 Tan,ＢＣ＆Jia,Ｙ、１９９８．Acladisticsmdyoffamily Tsubota4773（HIRO)，ｔｈｉｓｓｍｄｙ；Lembophyllaceae SematophyllaceaeinChinaJ・HattoriBot・Lab， Doﾉﾉc力o〃"j叩Ｌｓｍｅ応j/b"伽（Mitt.）Ｎｏｇ，ＡＢ ８４:49-55. Ｏ19465,Japan，Honshu，HyogoPref，Oya-cho，Ten- ＆’999．Apreliminaryrevisionof taki，Ｊｕｎｅ27,1998,HAkiyamal4252（HYo),this 亀 ChineseSematophyllaceae、JHattoriBotLab86： smdy;Sematophyllaceae〃伽ＣＯ"c､ﾉ(/bﾉﾉα(Cardot） l-70 Ｈ・ＡＣｒｕｍ,ＡＢＯ95271,Japan,Kyushu,Kagoshima-ken， Tsubota，Ｈ､，Nakao，Ｎ､，Arikawa，Ｔ，Yamaguchi，Ｔ， YakushimalsL，Shiratani-unsuikyoGorge,600-900m Higuchi,Ｍ,DegUchi,Ｈ､＆Seki,Ｔ・’999．Apre- alt.,Marcｈ24,2002,HDegUchi36051（HIRO),this liminaryphylogenyofHypnales(Musci)asinferred study；HypnaceaeGo"α"ｊαノwgj"osa（Mitt.）Broth， fi･omchloroplastr6cLsequencedata・BIyoLRes、 ＡＢＯ94341,Japan,Honshu,Nagano-ken,Shimo-ina-gun， ７:233-248 Minami-shinano-mura,KajitaniRiver,670-690ｍａｌｔ・’ ，Yamaguchi,Ｔ，Seki，Ｔ＆Deguchi， ０ct」9,2000,TArikawa2855(TNS),thisstudy;ＣＯﾉｰ Ｈ、2000．Preliminaryphylogeneticrelationshipsof ﾉα"ｊａｓｐ/e"｡e"ｓ（Iishiba）Ｎｏｇ，ABO94340,Japan， thegenusBro/〃ｅＭ/αanditsalliedgenera（Hyp‐ Honshu，Chiba-ken，Sanbu-gunNarutou-machi，Ｃａ、 nales,Musci)basedonchloroplastr6cLsequence ５ｍａｌｔ.,Dec、1,2000,colLTFuruki,inherbMHigu- dataJ・HattoriBot､Lab､８８:79-99 chi39333（TNS),thissmdy;Climaciaceaedj"、c伽１ －，Akiyama，Ｈ､，Yamaguchi，Ｔ＆Deguchi，Ｈ、 deノルojm（Hedw.）ＦＷｅｂｅｒ＆DMohr,ABO19442, 2001aMolecularphylogenyoftheSematophyl- Japan,Honshu,MiePref,Nabari-shLNagaki,March laceae(Hypnales,Musci)basedonchloroplastlbcL l6，1998,HAkiyamal4150（HYo),thissmdy;Ｃ〃 sequences．Ｊ・HattoriBotLab90:２２１－２４０． "１αc川zﾉ叩o"jc""ZLindh,ABO19443,Japan,Honshu， ＆２００ｌｂ・Molecular NaganoPrefljima-cho,ShiOjidaira,July28,1996,H phylogenyofthegenus刀jsmegjsrjaandrelated Akiyamal4376（HYo),ｔｈｉｓｓｍｄｙ;NeckeraceaeBパー genera(SematophyllaceaaMusci)basedonchloro- Ｓｃﾉﾉαﾉﾉ"g"〃α(Mitt.)Broth,ABO94789,Japan,Hon- plastrbcLsequencesHikobial3:529-549． shu，ＮａｒａＰｒｅｆ，Kamikitayama-mura，Odaigahara， Vanderpoorten,Ａ､,Hedeniis,Ｌ,ＣＯＸ,Ｃｌ＆Ｓｈａｗ,Ａ， l450malt.,Ｍａｙ26,1998,HAkiyamal4195(HYo)， J2002Circumscription，classification，ａｎｄtax- thisstudy;ﾊﾉO"!αﾉﾉαc/blIphzJs/αZgjo"/α""s(Mitt.)Dixon onomyofAmblystegiaceae（Bryopsida）ｉｎferred ＆ＰｄｅｌａＶａｒｄｅ，ABO94792,Japan，Shikoku，Kochi １ fromnuclearandchloroplastDNAsequencedata Pref，Monobe-mura，Befilkyo，９５０ｍａｌｔ.，Ｊｕｌｙ27, andmorphology・Ｔａｘｏｎ５１：ｌｌ５－ｌ２２ 1998,HAkiyamal4359(HYo),thisstudy;ｈ、α/jo- Ｖｉｔｔ,，.Ｈ、１９８４．ClassificationoftheBIyopsida・Ｉ､： `/e"｡〉℃〃scallpeﾉﾉ(/b/伽(Mitt.)MFleischABO94788， ⑨ RMSchuster（ed.),NewManualofBIyology， Japan，Honshu，HyogoPref，Kamigori-chq ＶＯＬ２，ｐｐ､696-759．HattoriBot・Ｌａｂ,Nichinan Onaru-keikoku,２５０ｍａｌｔ.,Ｊｕｎｅ１７，１９９８，HAkiyama l4234（HYo)，ｔｈｉｓstudy;ノVecAcrqp伽〃ｊ/ｊｑＭａ（Mitt.） AppendixA・Infbnnationonspecimensfiomwhich MFleisch，ＡＢＯ94790,Japan，Honshu，Okayama DNAwasextractedinthissmdy・ PrefNishiawakura-son,WakasugiPrimaryForest, Hereisalistoftaxawhoser6cLsequenceswereuti- ＬＯＯＯｍａｌｔ.,June17,1998,HAkiyamal4210(HYo)， lizedinthissmdywiththeiraccessionnumbers,show- thisstudy;Ｐｊ""areノルα"16秒α(Bosch＆SandeLac.） ingtheirsourcesandvoucherspecimeninfbnnation MF1eisch,ＡＢＯ94787,Taiwan,Pingung,MtPeitawu， VoucherspecimensarekeptinHIRqHYo，ｏｒＴＮｓ・ ｌ９５０ｍａｌｔ.,August２４，１９８８，Ｈ・AkiyamaTaiwan-l77 TreatmentoffhmiliesfbllowsVitt（1984）andBuck (HYo),thisstudy;Ｔｈａｍ"o6ﾉＷｍｓ"6ｓｅ７ｉａｊ""z(Mittex andVitt(1986） SandeLac.）ＢＣＴan，ABO94791［ａｓ７１ｈａｍ"o6ﾉＷｍ Thesequenceofinibrmationis：ｔｈｅｎａｍｅｏｆｔａｘｏｎ， ｓα"dejlJapan，Honshu,HyogoPref，Kamigori-cho， Ｈ・TsuBoTA,ＴＡＲＩＫＡｗＡ,Ｈ､ＡＫＩＹＡＭＡ,ＥＤＥＬｕＮＡ,Ｄ､GoNzALEz,MHIGucHlANDH､DEGucHI ６６５ Onam-keikoku,250malt.,Junel7,l99LH・Akiyama Scleanandourprogramsdownloadsite l423］（HYo)，ｔｈｉｓｓｍｄｙ；LeucodontaceaeLezJcM)〃 ＜http://homehiroshima-uacjp/chubo/＞ scj"Ｍ伽(Hedw）SchwiigF.,ABO95988,Japan,Hon‐ Ｄα/αbQse shu,NiigataPref,Itoigawa-shi,Kotaki,300ｍａｌｔ.,Ｊｕｌｙ DDBJ＜http://www・ddbjnigacjp/〉 28,1994,HAkiyamal2594(HYo),thisstudyMyu- Sequencedatadownloadsite 鰯 (HYo),thisstudy;Pterobryaceae伽c/(〕ﾉﾉollTqj"`/ｊｃｚｌ'〃 坪田博美･有川智己･秋山弘之･ElTainDeLuna･DDlores Mitt.,ＡＢＯ19464,Taiwan,Pingung,MtPeitawu，ｌ９５０ Gonzalez･樋口正信・出口博則：葉緑体ｒ６ｃＬ遺伝子の ｍａｌｔ.,HAkiyamaTaiwan-50(HYo),thisstudy． 大データセットに基づく腋蘇類とくにハイゴケ科とその 公 周辺科の分子系統学的研究 AppendixBInternetresourcesfbrthephylogenetic analysisanddatabase ハイゴケロ･イタチゴケ目･アブラゴケ目を含む腋蘇 SMw”e 類の系統関係を,葉緑体ｍｃＬ遺伝子の１８１の配列を含 CONSEL＜http://wwwjsmacjp/~shimo/prog/consel／ indexｈｔｍｌ＞ む大データセットに基づいて解析を行った．解析の結 MOLPHY＜fip://fipjsmacjp/pub/ISMLIB/MOLPHY/〉 果，ハイゴケ目とイタチゴケ目が－つの大きなクレー ＰＡＵＰ＊＜http://paupcsit・fSu・edu/＞ ドを形成したDeLuna他(2000)およびBuck他(2000） PAUPRat＜http://viceroy・eehuconn・edu/paupratweb／ の結果を支持したが，Vitt（1984）のハイゴケ目および pauprathtm＞ イタチゴケ目それぞれの単系統性は支持されなかっ TREE-PUZZLE＜http://www､tree-puzzlede/＞ た． " の